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South Atlantic Zooplankton

Vol. 2 Pages 869 - 1706

Edited by Demetrio Boltovskoy

Backhuys Publishers, Leiden 1999

Larval Decapoda (Brachyura) Gerhard Pohle, Fernando L. M Mantelatto, Maria L. Negreiros-Fransozo and Adilson Fransozo

Introduction

A proper knowledge of zooplankton, including its larval members, is of fundamental importance since it forms a vital link between primary producers and different consumer levels in the food chain (Wickstead, 1976). Crustacea are among the most prominent animals of the marine zooplankton (Hardy, 1958), to which they often contribute 50% or more of the bio-mass (Friedrich, 1969a). During their life cycles most decapod crustaceans spend some time as part of this community. With about 10,000 species (Bowman and Abele, 1982), the "ten-footed" Decapoda (Greek deka: ten, and pous: foot) represent the largest and most varied order of crustaceans, encompassing about one-third of known crustacean species. This order includes the typical larger and well known crustaceans, many of which live on or close to the bottom of the sea as juveniles and adults, but spend their larval life as part of the plankton.

Decapods have two basic adult body plans. Shrimps and lobsters possess well developed tail sections. In contrast, the Brachyura, or short-tailed decapods, have a flat abdomen flexed under the body. This group comprises the true crabs. With about 5,000 species worldwide (Melo, 1996), the true crabs represent half of the Decapoda. Crabs reach their greatest diversity in tropical regions, although a significant number are also found in temperate waters. Thus, within the South Atlantic alone, 328 crab species are presently recognized that belong to 170 genera among 24 families (Table 1). The recent accounts of Boschi et ah (1992), Zolessi and Philippi (1995), Martins and DTncao (1996), and Melo (1996) include most of these species.

Perhaps surprisingly, most of the larvae of these crabs remain unknown. Larval information presented here is available for 102 species, representing less than one third of all known crab species within the South Atlantic. While far from complete, this denotes significant progress over the last 16 years. In comparison, the last coverage of decapod larvae from the area (Boschi, 1981) included only about one quarter of the brachyuran species covered here. Nevertheless, it is

clear that we presently still have limited ability in identifying the decapod larvae from the South Atlantic. For example, larvae of several families, including the Cymonomidae, Raninidae, Cyclodorippidae, Geryoni-dae, Goneplacidae and Palicidae, are unknown within the South Atlantic (Table 1).

Among crustaceans, decapods are considered to be amongst the most advanced groups. Within the lineage of crawling decapods, or Reptantia (Latin reptare: to creep), the brachyurans represent the most evolutionary advanced forms, together with their sister group, the Anomala, comprising the hermit crabs and their relatives (Scholtz andRichter, 1995). Our understanding of the evolution of different brachyuran and other decapod groups, however, is still quite poor. Knowledge of ancestor-descendant relationships is largely based on adults. They display a vast array of highly specialized adaptations that may mask their evolutionary relationships. Evidence from larvae may help resolve these problems and early developmental stages are now increasingly used in phylogenetic reconstruction (Rice, 1980; 1983; Clark and Webber, 1991; Marques and Pohle, 1995; Pohle and Marques, 1998).

Even though larval types of most decapods are described below, the brachyuran larval stages found within the plankton are the focus of this chapter. During this phase of their life-cycle, larvae bear very little resemblance to the juvenile and adult form, and the inexperienced observer would be hard pressed to recognize the developmental stages of crabs. In fact, naturalists of an earlier day believed that such larvae represented different animals (Schmitt, 1971).

Before hatching, eggs of true crabs are extruded and brooded by the female in the space between the thorax and cupped abdomen. The number of eggs produced per brood varies widely, from as little as 200 (Telford, 1978), to as many as 8,000,000 (Prager et ah, 1990; Mantelatto and Fransozo, 1997). In tropical and subtropical areas, most species spawn and hatch batches of eggs throughout the year (Negreiros-Fransozo and Fransozo, 1995; Negreiros-Fransozo et ah, in press; Mantelatto and Fransozo, 1998). In these

South Atlantic Zooplankton. edited by D. Boltovskoy, pp. 1281-1351 © 1999 Backhuys Publishers, Leiden, The Netherlands

1282 Larval Decapoda (Brachyura)

regions incubation periods may be as short as 1-2 weeks (Pohle, 1994), egg size and temperature being determining factors. Larval development is also temperature dependent (Christiansen, 1973), a higher temperature shortening the period, and salinity also affects the duration of the larval phase (Fransozo and Negreiros-Fransozo, 1986; Negreiros-Fransozo and Fransozo, 1990). However, the number of larval stages is another determining factor in the length of the larval period. A warm-water species with 5 larval stages can reach the last larval stage in as little as 9-10 days after hatching (Marques and Pohle, 1996a; Fransozo and Negreiros-Fransozo, 1997).

Brachyurans have two distinct types of larvae, the zoea and megalopa. Zoeae emerge from eggs that usually hatch at night. In some species and under certain conditions, eggs may hatch as a prezoea before molting to a zoea. However, these short-lived prezoeal stages, that are still enclosed within a thin cuticle, are not usually found in the plankton.

Zoeae of various species appear very different from juvenile and adult stages (Fig. 2) but are themselves superficially alike. Thus all zoeae usually have large paired eyes and a full complement of carapace spines, consisting of a dorsal, rostral and lateral spines that give specimens a triangular upright appearance. A notable exception are dromiid and homolid larvae that look more shrimp-like (Fig. 4, 5). An abdomen consisting of a number of somites and ending in a flat fork-shaped telson protrudes from the carapace, as do two pairs of swimming appendages. Other carapace appendages are less apparent. The number of zoeal stages may vary from a single (Goodbody, 1960) to more than ten (Brossi-Garcia and Rodrigues, 1993; Cuesta and Rodrigues, 1994), depending on the species. Only within the family Majidae are there always only two zoeal stages. Older zoeal stages have the same general appearance but can be recognized by movable eyes, paired buds of appendages on the abdomen, rudiments of claws and legs under the carapace and by an increasing number of swirnrrifng setae on the locomotory appendages. However, for a considerable number of species only the first zoea is known.

The last zoeal stage undergoes a metamorphosis during the molt to the megalopa. The latter also have been referred to as the megalops (Sastry, 1970), megalop (Clark et al, 1998) decapodid (Felder et ah,

1985) or postlarva (Gurney, 1942). This stage has a more flattened, crab-like appearance, with legs and claws protruding from the carapace (Fig. 3). The spines of the carapace have either been lost or are greatly reduced. Unlike adult crabs, however, the abdomen has appendages used for swimming when it is unfolded, with the attachments folded under the carapace while at rest. In comparison to other stages, larval information on the megalopa is the poorest because it is not described in a number of larval publications. This is likely associated with difficulties in rearing, metamorphosis to the megalopa resulting in high mortality. This terminal larval stage is a transitional stage that settles out of the plankton and molts into the first crab instar, the first fully benthic stage. Additional morphological details of larval stages are given in the section dealing with identification.

Brachyuran zoeae are only a few millimeters in size, but are vigorous swimmers. Using both the maxilli-peds and abdomen to propel themselves upward and forward in pulses, with the dorsal spine often pointing in the direction of swimming, they swim at speeds of about 1-2 cm s_1 (Warner, 1977). When not active, zoeae sink, and thus they must constantly swim upwards to remain in the same place. In contrast, the megalopa swims smoothly forward with the dorsal spine in a vertical position, using abdominal appendages for propulsion. During swimming the legs are tucked close to the body to minimize resistance. Like many larval forms, zoeae react positively to light and a megalopa is initially also attracted to light but this is no longer the case during settlement.

Zoeal stages will feed on a large variety of phyto- and zooplanktonic organisms, appropriate size being more of a determining factor than type of food. However, evidence suggests that animal food is essential to complete larval development (McConaugha, 1985). Zoeal stages use the abdomen in the capture and manipulation of food. The megalopa feeds on other decapod larvae, copepods and young fish, using the claws for prey seizure and holding.

For decapod larvae, as for all marine animals with planktonic larvae, there is high mortality during larval life. Survival to a newly settled crab has been estimated at less than one tenth of a percent (Warner, 1967) for a single brood. This is compensated for by producing vast numbers of larvae. The advantage of produc-

Larval Decapoda (Brachyura) 1283

ing planktonic larvae is enhanced dispersal, allowing for rapid colonization in distant areas, and the general great abundance of food in the plankton.

Given the present limited knowledge of South Atlantic larvae, it is important for the reader to realize that larvae being identified may not fit any of those described herein. However, in such cases the keys, tables and figures will help in narrowing the search to higher groups, such as families. It also must be recognized that most of the larval accounts are based on laboratory rearings, and it is still unclear how much variability there is between specimens obtained from the wild and those obtained from culture (Ingle, 1992). Thus definitive identifications should be obtained by consultations with experts in the field.

For further reading on decapod larvae and their development, the reviews of Gurney (1942), Rice (1980), Williamson (1982) and Gore (1985) are recommended, as well as other references summarized at the end of this section.

Methods

Field collection

Crustacean larvae, like most other planktonic organisms, are collected in the wild using nets of various mesh sizes that are generally towed behind a vessel or left drifting where there are sufficient currents. Mesh sizes of 500 um are generally sufficient. A 4-5% formaldehyde (10:1 dilution of commercial formalin) solution is adequate for fixation of specimens. However, for final preservation and handling in the laboratory, transfer to 70% ethanol or 50% isopropanol is recommended.

Laboratory rearing

Undescribed larvae can only be positively identified when raised from known parentage. The first zoeal stage may be obtained by keeping a berried female until hatching occurs. However, other developmental stages can only be acquired by raising larvae in the laboratory. The simple tackle box (Costlow and Bookhout, 1959; 1960a) and finger bowl (Costlow and Bookhout, 1960b) techniques still suffice today. This includes regular changes of sea water and feed

ing with freshly hatched nauplii of the brine shrimp Artemia and the rotiferan Brachionus, readily available commercially in the form of dry cysts and resting eggs, respectively. This food source will suffice for larvae of many, but not all species. For those interested in more details of decapod larval culturing techniques, mass culture and single rearing methods have been examined and developed by Provenzano (1967), Rice and Williamson (1970), Sastry (1970), Roberts (1975), Kinne (1977) and Dawirs (1982).

Examination

A good stereo dissecting microscope is essential for proper examination of specimens that are no larger than a few mm in size. Both transmitted and reflected light should be used as a source of illumination, and a microscope equipped with a darkfield/brightfield base helps to highlight structural details.

Identification

For the purposes of this chapter, only the first zoea and the megalopa arc included. However, a key to staging of zoeae is provided. In general, the identifications are largely based on gross morphological features and relative size, and mostly only figures of the whole animal are presented. In some cases, however, this may not be satisfactory for proper identification.

The reader must also bear in mind that for some species not all larval stages are known and that for others, larvae are totally unknown. As well, it should be kept in mind that most descriptions are based on laboratory rearing and that all species exhibit some morphological variability. Thus, descriptions and illustrations may deviate from the specimens being compared. Consequently, when scientific reliability is essential, identifications of specimens should be verified by consulting with experts in the field or by cross-checking with the original description using the appropriate references cited. This may in many cases necessitate the dissection of appendages from the body to help in the identification. A brief protocol follows for anyone wishing to pursue this route.

Dissection will necessitate the use of very fine-tipped needles. While the finest insect pins may be satisfactory for relatively large specimens, best results are obtained by using tungsten wire that is electrolytical-


ly sharpened in a 10% KC1 solution and fastened onto a probe. For this a microscope transformer can be conveniently used, with one cable attached to an electrode (e.g. a nail) immersed in the solution, the other cable with the tungsten wire at the tip also being dipped into the liquid as low voltage is applied from the transformer. The voltage, emersion time, and depth of wire dipped, are manipulated until the desired shape is obtained.

With the specimen in water on a depression microscope slide, the larval carapace and abdomen should be separated before dealing with the appendages. This is done by using one needle to hold the carapace in place while gently pushing the abdomen away at its point of insertion until it detaches from the carapace. By piercing the carapace with one needle, the other needle can then be used to separate appendages at their point of attachment, starting from the posterior end of the larva. A stain, such as Chlorazol Black, can be added to increase contrast when necessary. For temporary mounts, the dissected parts are pipetted onto a flat microscope slide and a coverslip is applied. A sealant, such as clear fingernail polish, can be applied to the edges to help prevent evaporation. For the preparation of more permanent mounts the reader is referred to Koomen and Von Vaupel Klein (1995). A good compound microscope with lOx, 40x and lOOx objectives is required to examine the dissected appendages. A microscope equipped with phase contrast or Nomarski differential interference contrast would be preferable in order to facilitate the determination of critical features such as hair-like setae.

Geographic and bathymetric distribution

Known abiotic factors that strongly affect spatial distribution patterns of planktonic organisms include temperature, salinity, dissolved oxygen, water currents, and depth. The individual or collective action of these and other environmental factors increases or limits the area of distribution for marine species, where those factors with the most significant variations are the ones that limit the area of occurrence (Vernberg and Vernberg, 1970; Melo, 1985).

Presently the study of geographic and bathymetric distributions of larval crustacean stages is still in its infancy and the words of Gardiner (1904) that "... in

the present stage of knowledge any consideration of larval distribution is premature and must be inconclusive" still hold true today. The precise zoogeographi-cal distribution of many crab species that occur in the South Atlantic is especially poorly known in eastern waters. Larvae of many Southeastern Atlantic species are unknown and only those that also occur in western areas are covered in this volume. Distributional data presented are mainly based on the occurrence of adults in the Southwest Atlantic, along the coast of Brazil, Uruguay and Argentina. The Brazilian coast is the most diverse with approximately three hundred crab species (Table 1).

Taxonomy

Glossary

Italicized words are defined elsewhere in the glossary. PL: plural; abbr.: abbreviation.

Abdomen: the segmented hindmost part of the body usually consisting of 5-6 somites and telson.

Aesthetasc: specialized chemosensory seta with a thin cuticle; found on the antennule.

Antenna (pi. antennae; abbr.: an2): the second pair of segmented cephalic sensory appendages in Crustacea.

Antennule (abbr.: anl): the first pair of cephalic appendages.

Basis (pi. bases): the second segment of an appendage attached to the body.

Biramous: two-branched. Carapace: exoskeleton covering of cephalothorax,

often produced into dorsal, rostral and lateral spines. Cephalic: pertaining to the head. Cephalothorax: fused head and thorax (trunk). Cheliped: first pair of pereopods (ninth pair of

appendages), usually stouter than other pereopods, the last two segments forming a claw.

Coxa (pi. coxae): the basal segment of an appendage attached to the body.

Dactyl: the terminal or seventh segment of an appendage.

Denticulate: with small teeth. Endite: a non-specific term to describe a branch of an

appendage. Endopod: the inner branch of a biramous appendage. Epipod: lateral process attached to protopod.


S p e c i e s D i s t r i b u t i o n

Geographic Vertical (depth, m)

FAMILY DROMMDAE

Cryptodromiopsis antillensis (Stimpson, 1858)

Dromia erythropus (G. Edwards, 1771)

Hypoconcha arcuata Stimpson, 1858

Hypoconcha sabulosa (Linnaeus, 1 763) ( = H.

parasitica)

OC: North Carolina, Bermuda, Florida, Gulf of Mexico, The

West Indies, northern South America, Guianas and Brazil

(Amapa to Rio Grande do Sul)

OC: Bermuda, Florida, Gulf of Mexico, The West indies,

northern South America and Brazil (Pernambuco to Sao

Paulo) OC: North Carolina to Florida, Gulf of Mexico, The West

Indies, Guianas and Brazil (Amapa to Sao Paulo)

OC: North Carolina to Gulf of Mexico, The West Indies,

Venezuela and Brazil (Maranhao to Sao Paulo)

IT -330

SU - 360

S U - 8 0

SU - 9 0

FAMILY HOMOLI DAE

Homola barbata (Fabricius, 1793)

Thelxiope barbata (Fabricius, 1793)

OC: Virginia to South of Florida, Gulf of Mexico, The West

Indies, Central America, northern South America and Brazil

(Rio de Janeiro to Rio Grande do Sul); OR: Portugal and

Africa; Mediterranean Sea

OC: USA to Caribbean Sea, Brazil and Uruguay

(Maldonado); OR: Azores and Madeira Island, South Africa;

Mediterranean Sea

30 - 680

?

FAMILY LATREILLIIDAE

Latreiilia elegans Roux, 1828

Latreillia williamsi Melo, 1990

OC: North Atlantic, Brazil (Rio Grande do Sul) and

Uruguay; OR: Mediterranean Sea; Adriatic Sea

OC: Brazil (Rio de Janeiro to Rio Grande do Sul)

?

130-290; I

occasionally in

shallower

water

FAMILY DORIPPIDAE

Ethusa americana A. Milne Edwards, 1 880

Ethusa microphthaima Smith, 1881

Ethusa tenuipes Rathbun, 1897

Ethusina abyssicola Smith, 1884

OC: North Carolina, Florida, Gulf of Mexico, The West

Indies and Brazil (Maranhao to Rio de Janeiro); OP: Gulf of

California and Panama

OC: Massachussets to North Carolina, Florida, Gulf of

Mexico, The West Indies and Brazil (Sao Paulo)

OC: North Carolina, Florida, Gulf of Mexico, northern

South America and Brazil (Rio de Janeiro and Sao Paulo)

OC: Massachussets to North Carolina, Gulf of Mexico and

Brazil (Rio de Janeiro); OR: Mediterranean Sea: Spain

S U - 9 0

110- 750

40 - 400

850-4050

FAMILY CALAPPIDAE

Acanthocarpus alexandri Stimpson, 1871

Calappa angusta A. Milne Edwards, 1 880

Calappa gallus (Herbst, 1803)

OC: Massachussets, North Carolina to Florida, Gulf of

Mexico and Brazil (Rio de Janeiro to Rio Grande do Sul)


Indies, Venezuela and Brazil (Paraiba to Rio Grande do Sul)

OC: Bermuda, Florida, Gulf of Mexico, The West Indies,

Central America, northern South America and Brazil (Ceara

to Rio Grande do Sui); CA: Santa Helena Island; OR: Cape

Verde Island to Angola; Red Sea, Persian Gulf; IP: Japan

70 - 480

S U - 2 8 0

IT - 220




Calappa nitida Holthuis, 1958

Calappa ocellata Holthuis, 1958

Calappa sulcata Rathbun, 1898

Cycloes bairdii Stimpson, 1860

Hepatus gronovil Holthuis, 1959

Hepatus pudibundus (Herbst, 1 785)

Hepatus scaber Holthuis, 1959

Osachila antillensis Rathbun, 1916

Osachila tuberosa Stimpson, 1871

OC: The West Indies, Venezuela, Suriname, Guiana and

Brazil (Amapa and Para)


Indies, Colombia, Venezuela and Brazil (Amapa to Rio de

Janeiro)


Indies, Colombia, Venezuela, Guianas and Brazil (Amapato

Espirito Santo and Parana)


West Indies, Colombia, Venezuela and Brazil (Amapato Rio

de Janeiro); OP: California and Central America

OC: Colombia, Venezuela, Guianas and Brazil (Amapa to

Santa Catarina)

OC: Georgia, Gulf of Mexico, The West Indies, Venezuela,

Guianas, Brazil (Amapa to Rio Grande do Sul), Uruguay;

OR: Guinea to South Africa

OOVenezuela, Guianas and Brazil (Amapato Rio de

Janeiro)

OC: Bermuda, Gulf of Mexico, The West Indies and Brazil



Indies, Venezuela and Brazil (Rio de Janeiro to Rio Grande

do Sul)

S U - 7 0

S U - 8 0

SU -700

infralittoral -

230

2 0 - 6 0

SU - 1 60

2 0 - 8 5

80 - 300

4 0 - 1 9 0

FAMILY LEUCOSIIDAE

Callidactylus asper Stimpson, 1871

Ebalia stimpsoni A. Milne Edwards, 1880

lliacantha intermedia Miers, 1886

lliacantha liodactylus Rathbun, 1898

lliacantha sparsa Stimpson, 1871

lliacantha subglobosa Stimpson, 1871

* Leucosia planata (Fabricius, 1793)

Lithadia brasiliensis (von Martens, 1872)

Lithadia conica (Coelho, 1973)

Lithadia obliqua (Coelho, 1973)

Lithadia rotundata (A. Mi lne Edwards, 1880)

Lithadia vertiginosa (Coelho, 1973)

OC: North Carolina, Florida, Gulf of Mexico, Bermuda, The

West Indies, Colombia and Brazil (Amapa to Sergipe)


Indies, Colombia and Brazil (Amapato Sao Paulo)

OC: North and South Carolina, Florida, Guif of Mexico, The

West Indies, Colombia, Venezuela, Guianas and Brazil

i(ManinjTaoJ»J<!^^

OC: Florida, Gulf of Mexico, The West Indies, Colombia,

Venezuela, Guianas and Brazil (Amapato Bahia)

OC: Florida, Gulf of Mexico, The West Indies, Colombia and

Brazil (Para to EspTrito Santo)

OC: North Carolina to Florida, Gulf of Mexico, The West

Indies and Brazil (Amapa to Alagoas)

OC: Argentina

OC: Brazil (Para to Sao Paulo)

OC: Brazil (Amapa to Espfrito Santo)

OC: Brazil (Para to Pernambuco)

OC: Brazil (Rio Grande do Sul), Argentina (San Matias Gulf)

OC: Brazil (Para to Bahia)

2 5 - 9 0

S U - 1 6 0

SU - 130

S U - 1 3 0

2 0 - 8 0

SU - 400

?

S U - 4 0

4 0 - 8 5

S U - 3 0

2 0 - 8 0

3 0 - 6 0




Myropsis quinquespinosa Stimpson, 1871

Persephona crinita Rathbun, 1931

Persephona lichtensteinii Leach, 1 81 7

Persephona mediterranea (Herbst, 1794)

Persephona punctata (Linnaeus, 1758)

Randallia laevis (Borradaile, 1916)

Speloeophorus elevatus Rathbun, 1898

Speloeophorus nodosus (Bell, 1855)

OC: Massachussets, North Carolina, Florida, Gulf of Mexico,

The West Indies, Colombia, Venezuela, Guianas and Brazil

(Rio de Janeiro to Rio Grande do Sul), Uruguay and

Argentina

OC: Gulf of Mexico, The West Indies, Venezuela, and Brazil

(Amapa to Santa Catarina)

OC: Venezuela, Suriname, Guianas and Brazil (Amapa to

Sao Paulo)

OC: New Jersey, North and South Carolina, Florida, Gulf of

Mexico, The West indies, Venezuela, Suriname, Guianas

and Brazil (Amapa to Rio Grande do Sul) and Uruguay

OC: The West Indies, Colombia, Venezuela, Guianas and

Brazil (Amapa to Rio Grande do Sul)

OC: Brazil (llha Trindade)

OC: Florida, Gulf of Mexico, The West Indies and Brazil

(Maranhao to Bahia)

OC: North and South Carolina, Florida, Gulf of Mexico, The

West Indies and Brazil (Maranhao to Rio de Janeiro)

9 0 - 3 3 0

5 - 9 0

I T - 7 0

IT - 6 0

I T - 5 0

?

S U - 8 5

1 0 - 3 0

FAMILY CYCLODORIPP1DAE Clythrocerus carinatus Coelho, 1973

Clythrocerus granulatus (Rathbun, 1898)

Clythrocerus moreirai Tavares, 1 993

Cyclodorippe angulata Tavares, 1991

Cyclodorippe anfennan'a A. Milne Edwards, 1 880

Deilocerus analogus (Coelho, 1973)

Deiiocerus perpusilius (Rathbun, 1901)

Neocorycodusstimpsoni (Rathbun, 1937)


OC: Florida, The West Indies, Venezuela and Brazil (Amapa

to Rio Grande do Sul)

_0^_T3r^ZjMSJoJâulo) ,

OC: Brazil (Espfrito Santo and Rio de Janeiro)

OC: Gulf of Mexico, The West Indies and Brazil (Rio de

Janeiro)

OC: Brazil (Maranhao to Sao Paulo)

OC: North Carolina, The West Indies and Brazil (Amapa to

Rio Grande do Sul)


(Amapa to Sao Paulo)

2 0 - 6 0

80 - 600

65 - 220

130-350

40 - 650

60 - 110

3 0 - 1 8 0

4 0 - 1 8 0

FAMILY CYMONOM1DAE

Cymonomoides guinotae (Tavares, 1991)

Cymonomus guillei Tavares, 1991

Cymonomus magnirostris Tavares, 1991

Cymonomus quadratus A. Milne Edwards, 1880

OC: Brazil (Esplrito Santo and Rio de Janeiro)

OC: Brazil (Esplrito Santo and Rio de Janeiro)

OC: Brazil (Espirito Santo and Rio de Janeiro)


(Amapa and Rio de Janeiro to Rio Grande do Sul)

500 - 900

5 9 0 - 7 3 0

5 9 0 - 7 3 0

1 9 0 - 9 3 0

FAMILY RANINIDAE

Ranilia constricta (A. Milne Edwards, 1880)

Ranilia guinotae Melo and Campos Jr., 1994

Ranilia muricata H. Milne Edwards, 1837

Raninoides laevis (Latreille, 1825)

OC: Florida, Gulf of Mexico, The West Indies, and Brazil

(Amapa to Rio Grande do Sul); CA: Ascencao Island; OR:

Senegal to Congo

ÔcTBraz^KSôPeiulo)

OC: North Carolina to Gulf of Mexico, Colombia and Brazil

(Pernambuco)


Indies, Venezuela and Brazil (Amapa to Sao Paulo)

20 - 340

?

10 - 100

SU - 200



Geographic Vertical ; (depth, m) '

Symethis variolosa (Fabricius, 1793)


Indies and Brazil (Fernando de Noronha and Amapa to Sao

Paulo)

2 0 - 1 1 0 | I [

FAMILY MAj I DAE Acanthonyx dissimulatus Coelho, 1991-1993

Acanthonyx scutiformis (Dana, 1851)

Aepinus septemspinosus (A. Milne Edwards, 1879)

Anasimus fugax A. Milne Edwards, 1 880

Anasimus latus Rathbun, 1894

Anomalothir furcillatus (Stimpson, 1871)

Apiomithrax violaceus (A. Milne Edwards, 1868)

Arachnopsis filipes Stimpson, 1871

Batrachonotus brasiliensis Rathbun, 1894

Chorinus heros (Herbst, 1790)

Collodes armatus Rathbun, 1898

Collodes inermis A. Milne Edwards, 1878

Collodes rostratus A. Milne Edwards, 1878

Collodes trisplnosus Stimpson, 1871

Epialtoides rostratus Coelho, 1972

Epialtus bituberculatus H. Milne Edwards, 1834

Epialtus brasiliensis Dana, 1852

Euprognatha acuta A. Milne Edwards, 1 880

Euprognatha gracilipes A. Milne Edwards, 1878

Eurypodius latreillei Guerin, 1 828

Hemus cristulipes A. Milne Edwards, 1875

Herbstia depressa Stimpson, 1860

Holoplites armata (A. Milne Edwards, 1880)

OC: Brazil (Piaui to Bahia)

OC: Brazil (Espfrito Santo to Sao Paulo)


(Fernando de Noronha, Rocas and Para to Sao Paulo)

OC: The West Indies (Porto Rico) and Brazil (Amapa to Rio

de Janeiro)


Indies and Brazil (Amapa)


Indies and Brazil (Rio Grande do Sul)

OC: Brazil (Paraiba to Rio Grande do Sul); OR: Cape Verde

Island, Cape Branco to Angola; CA: Ascension Island


Indies and Brazil (Amapa and Rio Grande do Norte)



Venezuela and Brazil (Ceara to Bahia)

OC: Gulf of Mexico, Cuba and Brazil (Espfrito Santo and Rio

de Janeiro)

OC: Gulf of Mexico and The West Indies to Brazil (Amapa to

Rio de Janeiro)

OC: Brazil (Espfrito Santo to Rio Grande do Sul), Argentina

(including Patagonia)

OC: North Carolina to Florida, Gulf of Mexico and Brazil

(Amapa, Rio de Janeiro and Sao Paulo)

OC: Brazil (Maranhao to Espfrito Santo)


Venezuela and Brazil (Ceara to Sao Paulo)

OC: Colombia and Brazil (Espfrito Santo to Sao Paulo)

OC: Massachussets to Florida, Gulf of Mexico, The West

Indies, Guianas and Brazil (Amapa to Rio Grande do Sul)

and Uruguay


(Amapa and Ceara to Rio de Janeiro, Sao Paulo and Santa

Catarina)

OC: Brazil (Rio de Janeiro to Rio Grande do Sul), Argentina

(including Patagonia), Malvina Island, Strait of Magellan,

Uruguay; OP: Peru and Chile


West Indies and Brazil (Fernando de Noronha, Maranhao to

Rio de Janeiro)

OC: The West Indies, Venezuela and Brazil (Alagoas)

OC: Gulf of Mexico, The West Indies and Brazil (Para)

I T - 2 5 i

IT

1 0 - 8 5

60 - 200

S U - 1 6 0

50 - 1 80

1 0 - 5 0

30 - 240

1 2 - 7 3

1 0 - 5 0

2 0 - 7 0

S U - 4 0

2 0 - 6 5

1 0 - 2 5 0

2 0 - 6 0

IT

IT

1 5 - 7 1 0

70 - 370

S U - 1 0 0

1 5 - 70

60 - 700

1 60 - 800




Inachoides forceps A. Milne Edwards, 1879

Leptopisa setirostris (Stimpson, 1871)

Leucippa pentagona H. Milne Edwards, 1833

* Leurocyclusgracilipes (A. Milne Edwards and

Bouvier, 1923)

Leurocyclus tuberculosus (H. Milne Edwards and

Lucas, 1843)

Libidoclaea granaria H. Milne Edwards and Lucas,

1843

Libinia bellicosa Oliveira, 1944

Libinia ferreirae Brito Capello, 1871

Libinia spinosa H. Milne Edwards, 1834

Macrocoeloma camptocerum (Stimpson, 1871)

Macrocoeloma concavum Miers, 1886

Macrocoeloma eutheca (Stimpson, 1871)

Macrocoeloma laevigatum (Stimpson, 1860)

Macrocoeloma septemspinosum (Stimpson, 1 871)

Macrocoeloma subparallelum (Stimpson, 1 860)

Macrocoeloma trispinosum (Latreille, 1825)

Metoporhaphis calcarata (Say, 1818)

Microlissa brasiliensis (Rathbun, 1923)

Microphrys antillensis Rathbun,1920

Microphrys bicomutus (Latreille, 1 825)

Microphrys garthi (Lemos de Castro, 1953)

Microphrys interruptus Rathbun, 1920


Indies, Guianas and Brazil (Amapa to Rio de Janeiro)

OC: Florida, The West Indies, Venezuela and Brazil

(Maranhao to EspTrito Santo)

OC: Brazil (Rio de Janeiro and Sao Paulo), Uruguay and

Argentina; OP: California, Mexico and Chile

OC: Uruguay (Flores Island) and Argentina

OC: Brazil (Rio de Janeiro to Rio Grande do Sul), Uruguay

and Argentina (including Patagonia); OP: Chile

OC: Brazil (Rio Grande do Sul), Argentina (including

Patagonia) and Uruguay; OP: Chile (Strait of Magellan)

OC: Panama, Guianas and Brazil (Ceara, Rio de Janeiro and

Parana)

OC: Guianas, Venezuela, Brazil (Para to Santa Catarina) and

Uruguay

OC: Brazil (Espirito Santo to Rio Grande do Sul); OR:

Senegal to Angola, Cape Verde Island; OP: South of

California to North of Chile, Galapagos Islands and Hawaii

OC: North Carolina, Florida, Gulf of Mexico and Brazil

(Amapa to Maranhao)

OC: The West Indies and Brazil (Fernando de Noronha,

Maranhao to Bahia)


Indies, Central America and Brazil (Maranhao to Espfrito

Santo)


(Para to Alagoas)

OC: South Carolina, Florida, Gulf of Mexico and Brazil

(Ceara to Bahia)

OC: Gulf of Mexico, The West Indies, Venezuela and Brazil

(Fernando de Noronha, Amapa to Espfrito Santo)


West Indies and Brazil (Fernando de Noronha, Piauf to Sao

Paulo)


(Espirito Santo and Rio de Janeiro)

OC: Brazil (Ceara to Sao Paulo)


Indies and Brazil (Paraibato Rio de Janeiro)

OC: North Carolina to South of Florida, Bermuda, Gulf of

Mexico, The West Indies, Central America, Venezuela and

Brazil (Maranhao to Rio Grande do Sul; Fernando de

Noronha)

OC: Brazil (Parafbato Rio de Janeiro)

OC: The West Indies and Brazil (Piauf to Alagoas, Fernando

de Noronha)

1 5 - 7 0

I T - 8 0

2 0 - 8 0

?

10 - 1 70

deep

1 0 - 3 0

I T - 3 5

1 0 - 1 3 0

1 0 - 1 0 3

1 0 - 4 0

3 0 - 2 1 5

i T - 3 0

1 0 - 2 1 0

I T - 2 5

1 0 - 8 0

S U - 9 0

1 0 - 8 5

1 0 - 4 0

I T - 7 0

IT- 10

1 0 - 5 0




Mithraculus coryphe (Herbst, 1801)

Mithraculus forceps (A. Milne Edwards, 1875)

Mithraculus sculptus (Lamarck, 1818)

Mithrax besnardi Melo, 1990

Mithrax braziliensis Rathbun, 1892

Mithrax caribbaeus Rathbun, 1920

Mithrax hemphilli Rathbun, 1892

Mithrax hispidus (Herbst, 1790)

Mithrax tortugae Rathbun, 1920

Mithrax verrucosus H. Mi lne Edwards, 1832

Mocosoa crebripunctata Stimpson, 1871

Nemausa acuticornis (Stimpson, 1871)

Nemausa cornutus (Saussure, 1857)

Nibilia antilocapra (Stimpson, 1871)

Notolopas brasiliensis Miers, 1886

Paradasygius tuberculatus (Lemos de Castro, 1949)

Pelia rotunda A. Milne Edwards, 1875

Picroceroides tubularis Miers, 1886

Pitho Iherminieri (Schramm, 1867)

Podochela algicola (Stebbing, 1914)

Podochela brasiliensis Coelho, 1972

Podochela graciiipes Stimpson, 1871

Podochela minuscuia Coelho, 1972

OC: Florida, Gulf of Mexico, The West Indies, northern

South America and Brazil (Ceara to Sao Paulo; Fernando de

Noronha)

OC: North Carolina to South of Florida, Gulf of Mexico, The

West Indies, Venezuela and Brazil (Maranhao to Sao

Paulo; Rocas and Fernando de Noronha)

OC: Florida, Gulf of Mexico, The West Indies and Brazil (Rio

Grande do Norte to Bahia)

OC: Brazil (Rio Grande do Sul) and Uruguay

OC: Brazil (Piaui to Rio de Janeiro)

OC: The West Indies, Venezuela and Brazil (Bahia to Rio de

Janeiro)

OC: Florida, The West Indies and Brazil (Rocas and

Maranhao to Rio de Janeiro )

OC: Delaware to South of Florida, Gulf of Mexico, The West

Indies and Brazil (Para to Sao Paulo)

OC: Florida, The West Indies, Colombia, Venezuela and

Brazil (Espfrito Santo to Sao Paulo )

OC: South Carolina, Florida, Gulf of Mexico, The West

indies and Brazil (Fernando de Noronha and Rocas)

OC: Florida, Gulf of Mexico and Brazil (Maranhao to Rio de

Janeiro)


Indies and Brazil (Amapato Rio de Janeiro )

OC: Bermuda, Florida, Gulf of Mexico, The West Indies and

Brazil (Amapa to Bahia)


Indies and Brazil (Rio Grande do Norte to Rio Grande do

Sul)

OC: Colombia, Venezuela and Brazil (Amapato Sao Paulo)

OC: Brazil (Amapato Ceara)

OC: Brazil (Para to Rio Grande do Sul), Uruguay and

Argentina

OC: Florida, Gulf of Mexico, Cuba, Jamaica, Haiti, Saint

Thomas, Virgin Islands and Brazil (Maranhao to Espfrito

Santo)


Indies and Brazil (Para to Sao Paulo and Fernando de

Noronha)

OC: Colombia and Brazil (Maranhao to Sao Paulo)

OC: Brazil (Ceara to Sergipe)


West Indies, Colombia, Guianas and Brazil (Amapato Rio

Grande do Sul)

OC: Brazil (Ceara to Pernambuco)

I T - 6 0

I T - 9 0

10- '60

2 3 0 - 2 1 0 0

I T - 8

I T - 2 5

I T - 6 0

I T - 6 5

IT - 10

I T - 1 0

2 0 - 1 3 0

1 0 - 1 0 0

1 0 - 1 0 7 0

70 - 260

I T - 3 0

1 0 - 4 0

IT- 190

2 0 - 9 0

1 0 - 2 0 0

2 4 - 9 0

2 0 - 5 0

IT - 220

2 0 - 6 0




Podochela riisei Stimpson, 1860

Pyromaia tuberculata (Lockington, 1876)

Rochinia confusa Tavares, 1 991

Rochiniagracilipes A. Milne Edwards, 1875

Stenocionops furcata (Olivier, 1791)

Stenocionops spinimana (Rathbun, 1892)

Stenocionops spinosissima (Saussure, 1857)

Stenorhynchus seticornis (Herbst, 1788)

Taliepus dentatus (H. Milne Edwards, 1834)

Taliepus marginatus (Bell, 1835)

Teleophrys ornatus Rathbun, 1901

Teleophrys pococki Rathbun, 1924

Thoe aspera Rathbun, 1901

Tiche emarginata White, 1847

Tiche potiguara Garth, 1952

OC: North Carolina to Gulf of Mexico, The West Indies and

Brazil (Parafba, Pernambuco and Rio de Janeiro)

OC: Brazil (Rio de Janeiro to Parana); OP: California, Central

America to Chile; IP: Japan

OC: Brazil (Espirito Santo)

OC: Brazil (Rio de Janeiro to Rio Grande do Sul), Uruguay,

Argentina and Antarctica

OC: Georgia, Florida, Gulf of Mexico, The West Indies,

Colombia and Brazil (Ceara to Rio Grande do Sul)

OC: North Carolina to Florida, Gulf of Mexico and Brazil

(Sao Paulo)


Indies, Brazil (Fernando de Noronha, Rio Grande do Norte

to Rio Grande do Sul) and Uruguay

OC: North Carolina, Florida, Guif of Mexico, The West

Indies, Colombia, Venezuela (Isla Margarita), Guianas,

Brazil (Amapa to Rio Grande do Sul), Uruguay and

Argentina

OC: Brazil (Rio de Janeiro); OP: Peru and Chile

OC: Brazil and Uruguay; OP: Peru, Chile and Ecuador

(Galapagos Islands)

OC: Gulf of Mexico, The West Indies and Brazil (Fernando

de Noronha)

OC: Curacao and Brazil (Fernando de Noronha,

Pernambuco and Alagoas)

OC: Puerto Rico and Brazil (Pernambuco and Alagoas)


Indies and Brazil (Rio Grande do Norte)

OC: Brazil (Rio Grande do Norte to Alagoas)

S U - 1 4 0

1 0 - 1 3 0

5 9 0 - 7 3 0

1 5 - 1 7 5

IT -180

3 5 - 2 2 5

50 - 480

I T - 1 0 0

S U - 6 0

?

1 0 - 4 5

IT- 10

IT- 10

1 0 - 4 0

2 5 - 7 0

FAMILY PARTHENOPIDAE

Cryptopodia concava Stimpson, 1871

Heterocrypta granulata (Gibbes, 1 850)

Heterocrypta lapidea Rathbun, 1901

Heterocrypta tommasii Rodrigues da Costa, 1959

*Lambrus meridionalis Boschi, 1965

Leiolambrus nitidus Rathbun, 1901

Mesorhoea sexsp/nosa Stimpson, 1871

Parthenope (Parthenope) agona (Stimpson, 1871)

Parthenope (Piatylambrus) aylthoni (Righi, 1965)


Indies and Brazil (Maranhao to Rio de Janeiro)


Indies and Brazil (Ceara to Parana)

OC: The West Indies and Brazil (Para to Rio Grande do Sul)

OC: Central America, Guianas and Brazil (Ceara to Rio

Grande do Sul)

OC: USA (Pennsylvania) and Uruguay; Mediterranean and

Adriatic Sea; OR: Azores and Cape Verde Islands

OC: Gulf of Mexico, The West Indies, Guianas and Brazil

(Para to Espirito Santo)


Indies and Brazil (Para to Rio Grande do Sul)


Indies, Guianas and Brazil (Amapa to Rio Grande do Sul)

OC: Brazil (Rio de Janeiro and Sao Paulo), Uruguay and

Argentina

1 0 - 6 0

1 0 - 1 4 0

IT - 1 80

I T - 1 5

?

7 - 7 5

I T - 1 0 0

IT - 100

1 5 - 115



Geographic Vertical ] (depth, m) i

Parthenope (Piatylambrus) fraterculus (Stimpson,

1871)

Parthenope (Piatylambrus) guerini (Brito Capello,

1871)

Parthenope (Piatylambrus) pourtalesii (Stimpson,

1871)

Parthenope (Piatylambrus) serrata (H. Milne

Edwards, 1834)

Solenolambrus brasiliensis Rodrigues da Costa,

1961

Solenolambrus typicus Stimpson, 1871

Thyrolambrus astroides Rathbun, 1894


Indies, Suriname and Brazil (Amapa to Rio Grande do Sul)

OC: The West Indies and Brazil (Rio Grande do Norte to Sao

Paulo)

OC: New Jersey to South Florida, Gulf of Mexico, The West

Indies and Brazil (Amapa to Rio Grande do Sul)

OC: North Carolina, Florida, Bermuda, Gulf of Mexico, The

West Indies, Guianas, northern South America and Brazil

(Maranhao to Sao Paulo)

OC: Brazil (Rio de Janeiro to Santa Catarina)


Indies and Brazil (Rio de Janeiro)

OC: Gulf of Mexico, The West Indies and Brazil (Para to Rio

de Janeiro); IP: Mauritius and Andaman Sea; OP:

Revillagigedo Islands

1 0 - 2 0 0 |

1 5 - 3 0

20 - 350

1 0 - 1 1 0

1 0 - 100

9 0 - 6 2 0

5 0 - 3 7 0

FAMILY HYMENOSOMATIDAE

* Hallcarcinus planatus (Fabricius, 1775) OC: Argentina (Mar del Plata), Strait of Magellan; OP: Chile;

IP: New Zealand 10 - 1 70

FAMILY ATELECYCLIDAE

Peltarion splnulosum (White, 1843)

OC: Brazil (Rio Grande do Sul), Argentina (including

Patagonia) and Uruguay (Maldonado); OP: Chile (Punta

Arenas)

1 0 - 3 0 0

FAMILY BELLI I DAE

* Acanf/iocyc/us albatrossis Rathbun, 1898

* Acanthocyclusgayi Milne Edwards and Lucas,

1844

Bellia picta H. Milne Edwards, 1 848

Corystoides chilensis Lucas, 1 844

OC: Argentina (Malvinas Island); OP: Chile (Talcahuano and

Strait of Magellan)

OC: Argentina; OP: Chile

OC: Brazil (Rio Grande do Sul); OP: Peru and Chile

OC: Brazil (Rio de Janeiro to Rio Grande do Sul), Argentina

(Patagonia) and Uruguay; OP: Chile

SU

SU

?

5 - 3 0

FAMILY PORTUNI DAE

Arenaeus cribrarius (Lamarck, 1 818)

Calllnectes acutldens Rathbun, 1895

Callinectes bocourti A. Milne Edwards, 1879

Callinectes danae Smith, 1869

Callinectes exasperatus (Gerstaecker, 1856)

OC: Massachussets to North Carolina, Bermuda, Florida,

Gulf of Mexico, The West Indies, Colombia, Venezuela,

Brazil (Ceara to Rio Grande do Sul) and Uruguay

(Maldonado)

OC: Florida, Porto Rico, Panama, Brazil, Uruguay and

Argentina (Buenos Aires and Prata River)


Venezuela, Guianas and Brazil (Amapa to Santa Catarina)


Colombia, Venezuela and Brazil (Paraibato Rio Grande do

Sul)


Venezuela and Brazil (Maranhao to Santa Catarina)

I T - 7 0

?

I T - 2 0

I T - 7 5

I T - 8




Callinectes larvatus Ordway, 1863

Callinectes ornatus Ordway, 1863

Callinectes sapidus Rathbun, 1896

Charybdis hellerii (A. Mi lne Edwards, 1867)

Coenophthalmus tridentatus A. Milne Edwards,

1879

Cronius ruber (Lamarck, 1 818)

Cronius tumidulus (Stimpson, 1871)

Laleonectes vocans (A. Milne Edwards, 1878)

* Ovalipes catharus (White, 1 843)

Ovalipes punctatus (De Haan, 1833)

• Ovalipes trimaculatus (De Haan, 1833)

Portunus anceps (Saussure, 1 858)

Portunus gibbesii (Stimpson, 1859)

Portunus ordway] (Stimpson, 1860)

Portunus rufiremus Holthuis, 1959

OC: North Carolina to Florida, Gulf of Mexico, Bermuda,

The West Indies, Colombia, Venezuela and Brazil (Ceara to

Sao Paulo)


Indies, Colombia, Venezuela, Guianas and Brazil (Amapa to

Rio Grande do Sul)

OC: Southern USA to Gulf of Mexico, The West Indies,

Central America, Venezuela, Brazil (Bahia to Rio Grande do

Sul), Argentina and Uruguay; OR: North Sea, Mediterranean,

Adriatic Sea, Black Sea; IP: Japan

OC: eastern Florida, Cuba, Colombia, Venezuela and Brazil

(Alagoas, Sao Paulo, Santa Catarina); OR: Eastern

Mediterranean: Israel and Egypt; IP: Japan, Philipinnes, New

Caledonia, Australia, Hawaii, and throughout the Indian

Ocean, including the Red Sea


and Argentina (including Patagonia)

OC: North Carolina to South Florida, Gulf of Mexico, The

West Indies, Central America, northern South America,

Guianas and Brazil (Amapa to Rio Grande do Sul); OR:

Senegal to Angola; OP: California to Peru and Galapagos

Islands


Guianas and Brazil (Para to Sao Paulo)

OC: Gulf of Mexico, The West Indies and Brazil (Espfrito

Santo and Rio de Janeiro); CA: Ascension Island; OR:

Madeira, Cape Verde and Annobon Islands

OC: Uruguay (La Paloma, Cape Santa Maria) and Argentina

(Puerto Madryn); OR: South Africa; OP: Peru, Chile; IP:

Japan, China, Australia and New Zealand

OC: Brazil (Rio Grande do Sul), Uruguay (Rocha and Cape

de Santa Maria) and Argentina (Chubut); OR: South Africa;

OP: Peru and Chile; IP: Japan, China, Australia and New

Zealand

OC: Southern USA, Central America, Brazil (Sao Paulo to

Rio Grande do Sul), Uruguay and Argentina (including

Patagonia); OR: South Africa; South IP, including Australia

and New Zealand, and OP: Peru and Chile


West Indies and Brazil (Amapa to Rio de Janeiro)

OC: Massachussets to Florida, Gulf of Mexico, Venezuela,

Guianas and Brazil (Bahia)


Indies, Venezuela, Guianas and Brazil (Amapa to Rio

Grande do Sul, Fernando de Noronha)

OC: Guianas and Brazil (Amapa to Para)

I T - 2 5

4 - 7 5

I T - 9 0

I T - 5 1

1 5 - 5 0

IT -110

10-75

4 0 - 3 1 0

?

?

?

?

I T - 9 0

IT -110

2 0 - 4 5



Geographic Vertical (depth, m) j

Portunus spinicarpus (Stimpson, 1871)

Portunus spinimanus Latreille, 1819

Portunus ventralis (A. Milne Edwards, 1879)

Scylla serrata (Forskal, 1 775)


West Indies, Colombia, Venezuela, Guianas, Brazil (Amapa

to Rio Grande do Sul) and Uruguay (Maldonado)

OC: New Jersey to South Florida, Bermuda, Gulf of Mexico,

The West Indies, Venezuela, Guianas and Brazil

(Pernambuco to Rio Grande do Sul)

OC: Georgia, Florida, Gulf of Mexico, The West Indies,

Venezuela and Brazil (Rio Grande do Norte to Rio de

Janeiro)

OC: Brazil (Sao Paulo); OP: Mozambique and South Africa;

IP: Philipinnes and Japan

IT - 550

IT - 90

IT - 25

i IT -15 !

FAMILY GERYONIDAE

Chaceon notialis Manning and Holthuis, 1989

Chaceon ramosae Manning, Tavares and

Albuquerque, 1989

OC: Brazil (Cape Frio), Uruguay and Argentina; OP: Chile;

OR: Norway, Angola; IP: Madagascar, New Caledonia

OC: Brazil (Espfrito Santo to Sao Paulo)

? |

600-1200 |

FAMILY XANTHIDAE

Actae acantha (H. Milne Edwards, 1 834)

Allactaea lithostrota Williams, 1974

Banareia palmeri (Rathbun, 1 894)

Carpilius corallinus (Herbst, 1783)

Cataleptodius floridanus (Gibbes, 1850)

Domecia acanthophora acanthophora (Desbonne

and Schramm, 1867)

Edwardsium spinimanus (H. Milne Edwards, 1834)

Eriphia gonagra (Fabricius, 1781)

Euryozius sanguineus (Linnaeus, 1767)

Eurypanopeus abbreviatus (Stimpson, 1860)


(Fernando de Noronha, Rocas and Amapa to Pernambuco)


Indies, northern South America and Brazil (Rio de Janeiro to

Rio Grande do Sul)


South America and Brazil (Rocas, Para to Espfrito Santo)


Venezuela, Brazil (Ceara to Pernambuco; Fernando de

Noronha)

OC: Florida, Bermuda, Gulf of Mexico, The West Indies,

Central America, northern South America, Brazil (Ceara to

Rio Grande do Sul; Fernando de Noronha, Rocas); OR:

Africa (Guinea to Gabon)


West Indies, northern South America and Brazil (Sao Pedro

and Sao Paulo Reefs, Paraiba and Pernambuco)

OC: The West Indies, Guianas and Brazil (Ceara to Rio

Grande do Sul)

OC: North Carolina, Bermuda, Florida, Guif of Mexico, The

West Indies, Central America, northern South America and

Brazil (Para to Santa Catarina)

OC: Brazil (Para to Ceara and Sao Pedro and Sao Paulo

Reefs); OR: Madeira Island to Angola, Ascension and Santa

Helena islands


Indies, northern South America and Brazil (Ceara to Rio

Grande do Sui)

IT -25

| 50 - 640

1 0 - 1 5 0

I T - 2 0

IT -35

IT -145

1 5 - 5 5

I T - 5

2 0 - 120

I T - 5


1 S p e c i e s

' D i s t r i b u t i o n


Eurypanopeus depressus (Smith, 1869)

Eurypanopeus dissimilis (Benedict and Rathbun,

1891)

Eurytium Hmosum (Say, 1818)

Carthiope barbadensis (Rathbun, 1921)

Garthiope spinipes (A. Milne Edwards, 1880)

Clyptoxanthus vermiculatus (Lamarck, 1818)

Hexapanopeus angustifrons (Benedict and

Rathbun, 1891)

Hexapanopeus caribbaeus (Stimpson, 1871)

Hexapanopeus pais(ensis Rathbun, 1930

Hexapanopeus schmitti Rathbun, 1930

Leptodius floridanus (Gibbes, 1850)

Mefybia thalamita Stimpson, 1871

Menlppe nodifrons Stimpson, 1 859

Micropanope nuttingi (Rathbun, 1 898)

Micropanope pusllla A. Milne Edwards, 1880

Micropanope scu!ptipes Stimpson, 1871

Micropanope urlnator (A. Milne Edwards, 1881)

Panopeus americanus Saussure, 1857

Panopeus austrobesus Williams, 1983

Panopeus bermudensis Benedict and Rathbun,

1981

Panopeus harttii Smith, 1869

Panopeus herbstii H. Milne Edwards, 1834

OC: Massachusetts through Florida to southern Texas,

Dutch West Indies, Bermuda, Brazil (Paraiba) and Uruguay


(Pernambuco to Santa Catarina)


Central America, northern South America and Brazil (Para to

Santa Catarina)

OC: Florida, Gulf of Mexico, The West Indies, and Brazil

(Espfrito Santo)

OC: Bermuda, Florida, Gulf of Mexico, Venezuela and

Brazil (Amapa to Espfrito Santo)

OC: Venezuela, Guianas and Brazil (Espfrito Santo); OR:

Angola

OC: Massachussets to North Carolina, Florida, Gulf of

Mexico, The West Indies and Brazil (Pernambuco to Santa

Catarina)

OC: The West Indies, northern South America and Brazil

(Para to Rio Grande do Sul)

OC: South Carolina, Florida, Gulf of Mexico, Brazil (Para to

Santa Catarina) and Uruguay (Maldonado and Rocha)

OC: Brazil (Ceara to Santa Catarina) and Uruguay

OC: Florida, Bermuda to Brazil (Sao Paulo)

OC: Florida, Gulf of Mexico, The West indies, northern

South America and Brazil (Amapa to Sao Paulo)

OC: Florida, Gulf of Mexico, The West Indies, Central

America, northern South America, Guianas and Brazil

(Maranhao to Santa Catarina); OR: Cape Verde Island to

Angola


Indies and Brazil (Amapa to Sao Paulo)


(Para to Paraiba)


West Indies and Brazil (Amapa to Rio de Janeiro).

OC: North Carolina, Florida, Guff of Mexico, The West

Indies and Brazil (Para and Maranhao)


Venezuela and Brazil (Maranhao to Santa Catarina)

OC: Brazil (Rio de Janeiro to Rio Grande do Sul)


South America, Guianas and Brazil (Ceara to Santa

Catarina); OP*. Mexico to Peru

OC: Florida, Gulf of Mexico, The West indies and Brazil

(Maranhao to Sao Paulo); CA: Ascension Island

OC: Boston, Massachussets, Bermuda and Brazil (Santa

Catarina)

?

?

I T - 5

1 0 - 3 0

I T - 6 0

1 0 - 6 5

IT- 140

IT -55

I T - 5

IT -25

IT -15

10 - 200

IT- 10

10 - 180

3 0 - 3 1 0

1 0 - 3 1 0

1 50 - 460

IT -25

I T - 3 0

IT -15

IT -25

I T - 2 2




Panopeus lacustris Desbonne, 1867

*Panopeus margentus Will iams and Boschi, 1990

*Panopeus meridionalis Will iams, 1983

Panopeus occidental is Saussure, 1857

Paractaea rufopunctata nodosa (Stimpson, 1860)

Pilumnoides coelhoi Guinot and Macpherson,

1987

Pilumnoides hassleri A. Mi lne Edwards, 1880

Pilumnus caribbaeus Desbonne and Schramm,

1867

Pilumnus dasypodus Kingsley, 1879

Pilumnus diomedeae Rathbun, 1894

Pilumnus floridanus Stimpson, 1871

Pilumnus quoyi H. Milne Edwards, 1834

Pilumnus reticulatus Stimpson, 1860

Pilumnus spinosissimus Rathbun, 1 898

Platypodiella spectabilis (Herbst, 1794)

Platyxantbus crenulatus A. Milne Edwards, 1879

Platyxanthus patagonicus A. Milne Edwards, 1879

Tetraxantbus bidentatus (A. Milne Edwards,1880)

Tetraxantbus rathbunae Chace, 1939

Xanthodius denticulatus (White, 1847)

Xantbodius parvulus (Fabricius, 1 793) ^~~

OC: Bermuda, Florida, The West Indies, Colombia and

Brazil (Maranhao to Rio de Janeiro); OP: Hawaii

OC: Argentina (Mar del Plata)

OC: Uruguay (Montevideo) and Argentina (Plata River to

Mar del Plata)


Indies, Central America, northern South America, Guianas

and Brazil (Ceara to Santa Catarina)


Indies, northern South America, Brazil (Amapato Rio de

Janeiro) and Uruguay; CA: Ascension Island

OC: Brazil (Bahiato Santa Catarina)


and Argentina to Strait of Magellan


South America and Brazil (Parafba to Santa Catarina)


West Indies, northern South America and Brazil (Para to

Santa Catarina)

OC: Gulf of Mexico, The West Indies and Brazil (Amapa to

Rio Grande do Sul)

OC: North Carolina to Florida, Gulf of Mexico, Central

America, The West Indies, Venezuela and Brazil (Alagoas to

Bahia)

OC: Guianas and Brazil (Amapato Sao Paulo)

OC: The West Indies, Central America, northern South

America, Brazil (Para to Rio Grande do Sul), Uruguay and

Argentina


Grande do Norte to Santa Catarina)


Venezuela and Brazil (Fernando de Noronha, Trindade

island and Rio Grande do Norte to Rio de Janeiro)

OC: Brazil (Santa Catarina to Rio Grande do Sul), Uruguay,

Argentina (including Patagonia)

OC: Brazil (Rio Grande do Sul), Uruguay and Argentina


OC: North Carolina, Gulf of Mexico, Brazil (Cape Frio and

Santa Catarina) and Uruguay (Maldonado).


Indies and Brazil (Parafba to Rio Grande do Sul)


Venezuela and Brazil (Ceara to Bahia; Sao Pedro and Sao

Paulo Reefs)


Venezuela and Brazil (Rocas and Fernando de Noronha)

IT

SU

SU

I T - 2 0

IT - 220

1 0 - 3 0

?

I T - 5 5

I T - 3 0

24 - 340

?

IT - 100

I T - 7 5

5 - 2 0

5 - 1 5

SU

> 6 0

?

20 - 500

I T - 1 5

IT - 10




FAMILY GONEPLACIDAE

Acidops cessacii (A. Mi lne Edwards, 1878)

Bathyplax typhla A. Milne Edwards, 1880

Chasmocarcinus cylindricus Rathbun, 1901

Chasmocarcinus peresi Rodrigues da Costa, 1968

Chasmocarcinus rathbuni Bouvier, 1917

Chasmocarcinus typicus Rathbun, 1898

Cycloplax pinnotheroides Guinot, 1969

Cyrtoplax spinidentata (Benedict, 1892)

Eucratopsis crassimanus (Dana, 1852)

Euryplax nitida Stimpson, 1859

Frevillea hirsuta (Borradaile, 1916)

Nanoplax xanthiformis (A. Milne Edwards, 1880)

Neopilumnoplax americana (Rathbun, 1 898)

Panoplax depressa Stimpson, 1871

Pseudorhombila octodentata Rathbun, 1906

Pseudorhombila quadridentata (Latreille, 1828)

Speocarcinus carolinensis Stimpson, 1859

Speocarcinus meloi D'lncao and Gomes da Silva,

1992

OC: Brazil (Maranhao); OR: Africa: Cape Verde Islands,

Senegal and Saint Thome and Annobon Islands


Indies and Brazil (Pernambuco and Alagoas)

OC: Gulf of Mexico, The West Indies and Brazil (Rio de

Janeiro and Sao Paulo)

OC: Brazil (Para to Bahia)

OC: Brazil (Rio Grande do Sul)

OC: The West Indies, northern South America, Guianas and

Brazil (Rio de Janeiro to Rio Grande do Sul)

OC: Guianas and Brazil (Amapa and Para)

OC: The West Indies and Brazil (Pernambuco to Rio Grande

do Sul)


(Bahia to Rio Grande do Sui)


West Indies and Brazil (Piauf to Santa Catarina)




Indies, northern South America and Brazil (Amapa to Rio de

Janeiro)

OC: North Carolina, Georgia, Florida, Gulf of Mexico, Cuba

and Brazil (Espfrito Santo); IP: Arabic Sea


(Amapa to Pernambuco)

OC: The West Indies and Brazil (Rio de Janeiro to Rio

Grande do Sul)


(Bahia)


Indies and Brazil (Amapa, Sao Paulo and Rio Grande do Sul)

OC: Brazil (Rio Grande do Sul)

I T - 3 0

400 - 11 00

1 5 - 1 9 0 0

1 5 - 2 5

120

25 - 200

I T - 1 5

1 0 - 1 5 0

1 0 - 8 0

1 0 - 9 0

7 0 - 1 5 0

1 0 - 3 8 0

1 3 0 - 8 0 0

1 0 - 1 0 0

1 0 - 2 0 0

55

I T - 1 5 0

50 - 1 60

FAMILY PINNOTHERIDAE

Clypeasterophilus stebbingi (Rathbun, 1918)

Dissodactylus crinitichelis Moreira, 1901 = D.

encopei Rathbun, 1901

Fabia insularis (Melo, 1971)

Fabia sebastianensis Rodrigues da Costa, 1970

Parapinnixa bouvieri Rathbun, 1 918

OC: Florida, Gulf of Mexico and Brazil (Sao Paulo and Santa

Catarina)


Indies, northern South America, Brazil (Para to Rio Grande

do Sul) and Argentina

OC: Brazil (Rio de Janeiro to Rio Grande do Sul) and

Argentina

OC: Brazil (Sao Paulo)


Indies and Brazil (Amapa)

5 - 1 0

5 - 5 0

2 - 4 0

5 - 2 0

5 - 7 5




Parapinnixa hendersoni Rathbun, 1918

* Pinnaxodes chilensis (A. Milne Edwards, 1837)

Pinnaxodes tomentosus Ortmann, 1894

Pinnixa brevipollex Rathbun, 1 898 (= Pinnixa

rapax Bouvier, 191 7)

Pinnixa aidae Righi, 1967

Pinnixa chaetopterana Stimpson, 1 860

Pinnixa cristata Rathbun, 1900

Pinnixa patagoniensis Rathbun, 1918

Pinnixa sayana Stimpson, 1860 "•'"

Pinnotheres emiliai (Melo, 1971)

Pinnotheres garthi Fenucci, 1975

Tumidotheres maculatus (Say, 1818)

Zaops ostreum (Say, 1 81 7)

OC: Florida, Gulf of Mexico, The West Indies, Venezuela

and Brazil (Maranhaoto Espirito Santo)

OC: Argentina; OP: Equador to Chile

OC: Brazil


and Argentina

OC: Brazil (Alagoasto Sao Paulo)

OC: Massachusetts to North Carolina, Florida, Gulf of

Mexico and Brazil (Pernambuco to Rio Grande do Sul)

OC: North Carolina to Florida, Gulf of Mexico, Central

America and Brazil (Amapa, Pernambuco and Sao Paulo)


and Argentina (Gulf of San Matias)

OC: Massachusetts to North Carolina, Florida, Gulf of

Mexico and Brazil (Amapa to Rio Grande do Sul)

OC: Brazil (Rio de Janeiro and Rio Grande do Sul) and

Argentina (Mar de! Plata)

OC: Brazil (Rio Grande do Sul) and Argentina (Mar del Plata,

Gulfs of Necochea and Saint Mathias)

OC: Massachusetts to Florida, Gulf of Mexico, The West

Indies, Brazil (Alagoas to Santa Catarina), Uruguay and

Argentina

OC: Massachusetts to South of Florida, Gulf of Mexico, The

West Indies and Brazil (Pernambuco to Santa Catarina)

4 0 - 6 0

5 - 1 0

?

3 0 - 7 0

5 - 1 0

I T - 6 0

I T - 1 0

5 - 1 0

1 0 - 7 5

1 0 - 2 1

5 - 10

I T - 5 0

I T - 1 0

FAMILY GRAPSIDAE

Aratuspisonii (H. Milne Edwards, 1837)

Armases angustipes (Dana, 1852)

Armases benedicti (Rathbun, 1897)

Armases miersii (Rathbun, 1897)

Chasmagnathus granulata Dana, 1851

Cyclograpsus integer H. Milne Edwards, 1837

Cyrtograpsus angulatus Dana, 1851

Cyrtograpsus affinis (Dana, 1851)

Cyrtograpsus altimanus Rathbun, 1914


South America, Guianas and Brazil (Piauf to Sao Paulo); OP:

Nicaragua to Peru, Chile

OC: Mexico, The West Indies and Brazil (Ceara to Santa

Catarina)

OC: Florida, Gulf of Mexico, Venezuela, Guianas and

Brazil (Amapa and Para)

OC: Central America, northern South America, Brazil and

Uruguay (Montevideo: Ratas Island)


and Argentina

OC: Florida, Gulf of Mexico, Central America, northern

South America and Brazil (Ceara to Santa Catarina); OR:

Cape Verde Island to Senegal; IP


(Montevideo) and Argentina (Rawson, Chubut and

Patagonia); OP: Peru and Chile

OC: Brazil (Rio de Janeiro to Rio Grande do Sul) and

Argentina; OP: Peru and Chile

OC: Brazil (Rio Grande do Sul), Uruguay and Argentina


SM and MA

SM and MA

SM and MA

SM and MA

SM and MA

SM and MA

IT

IT

IT



Geographic Vertical [ (depth, m)

Euchirograpsus americanus A. Mi lne Edwards, 1

1880

Euchirograpsus kingsleyi (Miers, 1885) .

Geograpsus lividus (H. Milne Edwards, 1837)

Goniopsis cruentata (Latreille, 1803)

Grapsus grapsus (= G. adcensionis ) (Linnaeus,

1758)

Metasesarma rubripes (Rathbun, 1897)

Pachygrapsus corrugatus (von Martens, 1872)

Pachygrapsus gracilis (Saussure, 1858)

Pachygrapsus transversus (Gibbes, 1850)

Percnon gibbesi (H. Milne Edwards, 1853)

Plagusia depressa (Fabricius, 1775)

Planes cyaneus Dana, 1851

*Planes minutus (Linneo, 1758)

Sesarma crassipes Cano, 1889

Sesarma rectum Randall, 1840


West Indies, Colombia, Venezuela and Brazil (Rio Grande

do Sul)

OC: Brazil (Rio Grande do Sul); OR: South Africa


northern South America and Brazil (Rio de Janeiro to Rio

Grande do Sul); OR: Senegal to Angola and Cape Verde

islands; OP: California to northern Chile, Galapagos Islands

and Hawaii


Guianas and Brazil (Fernando de Noronha, Para to Santa

Catarina); OR: Senegal to Angola


Colombia, Venezuela and Brazil (Ceara to Espfrito Santo;

Fernando de Noronha, Rocas and Trindade); OP: California

to Chile and Galapagos Islands

OC: Central America, northern South America, Guianas,

Brazil (Ceara to Rio Grande do Sul), Uruguay (Montevideo)

and Argentina

OC: The West Indies and Brazil (Sao Pedro and Sao Paulo

Reefs)


Indies, northern South America, Guianas and Brazil

(Trindade Island, Ceara to Rio Grande do Sul); OR: Cape

Verde Island to Angola, Mediterranean Sea; OP: California

to Peru

OC: Bermuda, North Carolina to Florida, Gulf of Mexico,

The West Indies, northern South America, Brazil (Trindade

Island, Ceara to Rio Grande do Sul) and Uruguay; OR: Cape

Verde Island to Angola, Mediterranean Sea; OP: California

to Peru


West Indies and Brazil (Fernando de Noronha); OR: Azores

to South Africa; OP: California to Chile, Galapagos Islands

and Clipperton Island


West Indies and Brazil (Fernando de Noronha, Rocas, Sao

Pedro and Sao Paulo Reefs, Trindade Island, Ceara to Bahia);

OR: Azores and Madeira, Senegal to Angola

OC: Brazil (Rio Grande do Norte to Rio Grande do Sul),

Uruguay and Argentina; OR: Tropical Africa and Santa

Helena Island; IP and OP: California, Peru and Chile

OC: Uruguay; Pacific ocean

OC: Costa Rica and Brazil (Pernambuco and Bahia)

OC: Venezuela, Guianas and Brazil (Amapato Santa

Catarina)

3 0 - 5 1 0

3 0 - 3 2 0

IT

IT

IT

SM and MA

IT

IT

IT

IT

IT

I T - 1 5

?

SM and MA

SM and MA




FAMILY GECARCINIDAE

Cardisoma guanhumi Latreille, 1825

Gecarcinus lagostoma H. Milne Edwards, 1835


Colombia, Venezuela and Brazil (Ceara to Sao Paulo)

OC: Florida, The West Indies, Venezuela and Brazil

(Fernando de Noronha, Rocas and Trindade); CA: Ascension

Island

MA

MA

FAMILY OCYPODIDAE

Ocypode quadrata (Fabricius, 1 787)

Uca burgersi Holthuis, 1967

Uca cumuianta Crane, 1943

Uca leptodactyla Rathbun, 1898

Uca maracoani (Latreille, 1802-1803)

Uca mordax (Smith, 1870)

Uca rapax (Smith, 1 870)

Uca thayeri Rathbun, 1900

Uca uruguayensis Nobil i , 1901

Uca victoriana von Hagen, 1987

Uca vocator (Herbst, 1 804)

Ucides cordatus (Linnaeus, 1763)

OC: Florida, Bermuda, Gulf of Mexico, Central America,

The West Indies, northern South America, Guianas and

Brazil (Fernando de Noronha, Para to Rio Grande do Sul)


and Brazil (Maranhao to Sao Paulo)

OC: Central America, northern South America, Guianas and

Brazil (Para to Rio de Janeiro)


and Brazil (Maranhao to Santa Catarina)

OC: The West Indies, Venezuela, Guianas and Brazil

(Maranhao to Parana)

OC: Gulf of Mexico, Central America, northern South

America, Guianas and Brazil (Para to Sao Paulo)


and Brazil (Para to Santa Catarina)

OC: Florida, Gulf of Mexico, The West Indies, Guatemala,

Panama, Venezuela and Brazil (Maranhao to Santa Catarina)


and Argentina

OC: Brazil (Espfrito Santo)

OC: Gulf of Mexico, The West indies, Central America,

northern South America, Guianas and Brazil (Pernambuco to

Santa Catarina)

OC: Florida, Gulf of Mexico, The West Indies, Central

America, northern South America, Guianas and Brazil (Para

to Santa Catarina)

IT

MA

MA

MA

MA

MA

MA

MA

MA i

MA

MA

MA

FAMILY PALIC1DAE

Palicus acutifrons (A. Milne Edwards, 1880)

Palicus affinis A, Milne Edwards and Bouvier,

1899

Palicus alternatus Rathbun, 1 897

Palicus dentatus (A. Milne Edwards, 1880)

Palicus faxoni Rathbun, 1897

Palicus obesus (A. Milne Edwards, 1 880)

Palicus sica (A. Milne Edwards, 1880)

OC: Brazil (Bahia and Espfrito Santo)

OC: Florida, Gulf of Mexico, The West Indies, Guianas and

Brazil (Maranhao to Espfrito Santo)

OC: North Carolina, Florida, Gulf of Mexico and Brazil (Rio

de Janeiro to Rio Grande do Sul)


de Janeiro to Rio Grande do Sul)

OC: North Carolina to Florida, Gulf of Mexico, Yucatan and

Brazil (Rio Grande do Norte to Rio de Janeiro)

OC: Florida, Gulf of Mexico, Mexico and Brazil (Rio de

Janeiro to Rio Grande do Sul)



1 0 - 3 0

2 0 - 2 1 5

1 0 - 1 1 0

3 0 - 1 4 0

3 5 - 9 5

2 0 - 2 2 0

1 0 - 1 90


D i s t r i b u t i o n S p e c i e s


FAMILY CRYPTOCHIRIDAE

Opecarcinus hypostegus (Shaw and Hopkins,

1977)


(Pernambuco); CA: Ascension Island S U - 2 7

iTroglocarcinus corallicola Verril, 1908

OC: Bermuda, Florida, Gulf of Mexico, The West Indies and

Brazil (Maranhao to Bahia; Fernando de Noronha, Rocas);

CA: Ascension and Santa Helena Islands; OR.: Gabon, Saint

Tome and Annobon

S U - 7 5

Table 1. The geographic and bathymetric distribution of brachyuran species recorded from the South Atlantic, but excluding species that are endemic to the Southeast Atlantic. Species marked with an asterisk were not recorded along the Brazilian coast. The depth distribution corresponds to the minimum and maximum depth of occunence for the species. This list is based on Williams (1984), Boschi et al. (1992), Zolessi and Philippi (1995), Carmona and Conde (1996), Martins and D'lncao (1996), and Melo (1996). (OC: Occidental Atlantic; OR: Oriental Atlantic; CA: Central Atlantic; OP: Oriental Pacific; IP: Indo Pacific; IT: intertidal zone; SU: sublittoral zone; SM: salt marshes; MA: mangrove).

Exopod: the outer branch of a biramous appendage. Flagellum (pi. flagella): distal endopod portion of

megalopal antenna beyond the peduncle that consists of several segments.

Furca (pi. furcae): the forked end of the abdomen in zoeal stages.

Ischium (pi. ischia): the third most basal segment of an appendage, or first segment of endopod articulating with basis.

Mandible (abbr.: md): third pair of appendages; first pair of mouthparts.

Maxilla (abbr.-. mx2): fifth pair of appendages; third pair of mouthparts.

Maxillule (abbr.: mxl): fourth pair of appendages; second pair of mouthparts.

Maxillipeds (abbr.: mxpl, mxp2...): sixth to eighth pair of appendages; the three most posterior pairs of mouthparts, the first two of which are used for swimming in zoeal stages.

Palp: in the megalopa the distal 1-3 segmented portion of the maxilliped endopod and the segmented structure attached to the mandible.

Peduncle: the proximal non-branched segments of the antenna and antennule in the megalopa.

Pereopods: ninth to thirteenth pair of appendages; the chelipeds plus four pairs of walking legs.

Pleopods: fourteenth to eighteenth pair of appendages on the second to sixth abdominal somites.

Protopod: the proximal part of a crustacean limb that lacks lateral branches, in zoeae usually consisting of the coxa and basis which may be fused, or of

more segments in the megalopa (peduncle); especially here the basal part of the zoeal antenna from which arises the spinous process.

Ramus (pi. rami): a branch of any branched limb; thus a biramous limb is divided into exopodite and endopodite; a uniramous limb has only one branch.

Rostrum: anterior median extension of the carapace often forming a distinct spine or spines.

Rostral spine: see rostrum. Seta (pi. setae): a bristle, spine- or hair-like structure,

with basal socket, and produced as an extension of the cuticle; mostly on appendages; setal shaft may be smooth (simple seta), or feathery (plumose seta).

Scaphognathite: epipod of the maxilla which regulates water flow past the respiratory surface.

Somite: a segment of the body, not of the appendage. Spine: direct and continuous outgrowth of exoskele-

ton, forming a gross morphological projection. Spinous process: in zoeae an extention of the antennal

protopodite bearing small spines. Telson: the unpaired appendage on the hindmost

abdominal somite (see abdomen) which bears the anus.

Thorax: seventh to fourteenth somites; in Brachyura always fused with the six head somites to form a cephalothorax.

Uniramous: single-branched, as opposed to biramous or two-branched.

Uropod: paired appendage on sixth abdominal somite (=pleopod 5).


Group

Dendrobranchiata

Nauplius; Protozoea

Penaeoidea 5, 6 (8) 3

Sergestoidea 2 3

Pleocyemata

Stenopodidea !

Caridea ~

Scyllaridae, Palinuridae

Nephropidae, Polychelidae

Thalassinidea

Anomura

Brachyura

-

I -

-

-

-

-

Mysis

2-5

2

-

-

-

"

Zoea Phyllosoma

-

- -

1-9

many

-

-

- ! 6-15

2-many

2-many

2-many

-

-

-

"Eryoneicus"

-

-

-

-

-

3

-

-

-

Decapodid

few

many

1

1(2,3)

1 (puerulus)

1

1

l(glaucothoe)

1 -2 (megalopa) mostly 1

Table 2. Number of larval stages in each phase for different groups of decapod Crustacea (modified from Williamson, 1982).

Recognition of the main decapod larval forms

According to Williamson (1982) and Gore (1985), the first larvae produced by decapods usually consist of a free-swimming or motile planktonic form that hatches from the egg. These larvae hatch as different types, ranging from morphologically relatively simple forms termed nauplii, among decapods found in dendrobran-chiate shrimps, to more complex forms found in ple-ocyematids, termed prezoea, zoea, naupliosoma or phyllosoma. Subsequent stages are either a megalopa, glaucothoe, puerulus or decapodid (Table 2). The collective term decapodid is used here as by Williamson (1982) and Felder et al. (1985).

The distinction between each larval phase is mainly based on the appendages used for locomotion. The phases are: 1. Nauplius (several stages): It is characterized by a

single small median eye and propulsion is by means of three pairs of head appendages (antennae and mandibles). This type of larva belongs to members of the Dendrobranchiata.

2. Protozoea (three stages): This larva has a pair of compound eyes, an antenna that is segmented to the base, locomotion that is shared between the second antennae and exopods of the first two max-illipeds, and a telson with two blunt cylindrical rami. This kind of larva belongs to representatives of the Dendrobranchiata.

6.

Prezoea (one stage): It is the last embryonic stage, which precedes the first stage zoea; its free life is usually only a matter of minutes before molting again. This stage is totally enveloped by a cuticle and, unlike for other phases, the appendages do not have setae. Movement is limited to abdominal flexion. This kind of larva can appear in Brachyuran development. Zoea (several stages): It is similar to the protozoea, from which it differs by the means of locomotion. In early zoeae, propulsion is by means of exopods of maxillipeds and in some older zoeae also by exopods of pereopods. The pleopods can be present but are not natatory. Zoeae are found as older larvae of dendrobranchiates or as early larvae of pleocyemates. Mysis (several stages): With setose swimming exopods on the maxillipeds and pereopods; pleopods are present starting with the second stage but lack setae and are not yet functional. Locomotion is by exopods of the thoracopods. This larval form can be found among penaeideans after the zoeal phase. Phyllosoma (several stages): The body is dorsoven-trally flattened, anteriorly pear-shaped, and wider than long or as long as wide; third maxillipeds with or without exopod, abdomen one tenth to one fourth of total length, and locomotion is by cephalothora-cic maxillipeds with well developed and numerous natatory setae on the exopods. These larvae are found among scyllarid and palinurid lobsters.


7. "Eryoneicus" (several stages): Body not extremely compressed; eyes rarely covered by carapace; at least two pairs of functional thoracopods; antennal exopod unsegmented or segmented at tip only; abdomen one third of total length (excluding rostrum) (Nephropidae and Polychelidae). There is some debate if this larva is equivalent to a zoeal or post-larval phase (Gurney, 1942; Williamson, 1982).

8. Decapodid (usually one stage): This is the transi-. tional stage between the zoeal and juvenile growth

phase. The locomotory function has shifted to setose pleopods for swimming but pereopods for crawling are also present. 8.1. Megalopa: This term is usually used for bra-

chyuran and some anomuran decapodids. It looks like a small crab with an extended abdomen bearing ventral pairs of pulsating setose pleopods while swimming. The pereopods are also well developed, the first pair being chelipeds.

8.2. Glaucothoe: This usually refers to an anomuran pagurid decapodid. The first pereopod is chelated; the 5th pereopod and, sometimes also the 4th, are distinctly smaller than the other pereopods.

8.3. Puerulus: is usually used for a palinuroidean decapodid. It looks like a small spiny lobster.

8.4. Penaeoidean decapodid: The most characteristic feature is that swimming shifts during this stage from the thorax to the abdomen.

Key to recognize zoeal larvae of major decapod groups (Fig. 1)

1 Telson cylindrical in cross section 2 la Telson flattened 3 2 Telson with two blunt rami: protozoea (Dendro-

branchiata) (Fig. 1A, B) 2a Telson not produced into blunt rami 4 3 Telson furcated: zoea (Brachyura) (Fig. 1M) 3a Telson not furcated 5 4 Body extremely flattened dorsoventrally; fore-

body pear-shaped, wider than long or as long as wide; maxilliped 3 with or without exopod; abdomen one tenth to one fourth of total length: Phyllosoma (Scyllaridae and Palinuridae) (Fig. IF)

4a Body not extremely flattened; eyes rarely covered by carapace; at least two pairs of functional thoracopods with chelae; antennal exopod

unsegmented or segmented at tip only; abdomen one third of total length (excluding rostrum): Eryoneicus (Polychelidae) (Fig. 1G); zoea (Nephropoidea) (Fig. 1H)

5 Telson spatula-like, with first marginal process never as a spine; carapace without dorsal and lateral spines; antennal scale segmented: zoea (Caridea) (Fig. ID)

5a Telson subtriangular 6 6 Lateral margin of the telson serrate and rostral

spine well developed: zoea (Stenopodidea) (Fig. 1C)

6a Lateral margin of the telson smooth 7 7 Posterior border of the telson bearing a conspicu

ous median spine, abdominal somites 2-6 with prominent dorsal median spine: zoea (Thalassini-dea) (Fig. IE)

7a Carapace longer than wide; telson formula 7+7, the first lateral process spinous and the second one hair-like; antennal scale more or less developed: zoea (Anomura and Dromiacea) (Fig. II, K,L)

During the zoeal phase each larval stage is easily recognized by morphological features, particularly the long swimming (natatory) setae on maxilliped appendages and characteristics of the abdomen (Table 3).

General morphology of brachyuran larvae

Zoeal phase (Fig. 2)

The main body, or cephalothorax, of a zoea is covered by a carapace with free ventral and posterolateral margins that enclose the proximal parts of incipient thoracic limbs, gill buds and internal organs. A pair of large eyes are located anteriorly and several paired appendages are inserted ventrally. A segmented flexible abdomen terminating in a telson is attached to the cephalothorax. The cephalothorax and abdomen are pigmented but these chromatophores fade in preserved specimens and thus are not shown in figures. Due to the arrangement of prominent spines on the carapace in the majority of larvae, zoeae of species in the area have a more or less upright triangular carapace, but in some species (Dromiidae, Homolidae and Latreilliidae) zoeae have a shrimp-like carapace that is longer than wide. Typically, the carapace bears a rostral, dorsal and two lateral spines but other combinations are possible.


GENERAL DECAPOD LARVAL FORMS

© Palinura, Scyllaridae


GENERAL DECAPOD LARVAL FORMS

(§) Palinura, Polychelidae (J> Astacura, Nephropoidea

O Anomura, Porcellanidae

Anomura, Hippidae O

o Anomura, Paguridae

(J) Brachyura, Portunidae

O Anomura, Diogenidae

Fig. 1. General decapod larval forms. Sources, from Boschi (1981): A, B, K, M; from Gore (1970): I; from Hebling and Fransozo (1982): L; from Knight (1967): J (originally fig. 1 on p. 63); from Rodrigues (1976): E; from Gurney (1942): G, H; from Williamson (1976): C; from Williamson (1982): F.


PHASE/STAGE

ZOEAI

ZOEA II

ZOEAII

FEATURES

ZOEA IV (and older stages)

DECAPODID (MEGALOPA)

Eyes sessile; antennule and antenna without endopod buds; first two maxillipeds natatory, exopod segmented and distally with 4 long setae; third maxiliiped absent or not visible; no uropods; telson not separated from 6th abdominal somite, groups with typical forked telson usually with 3 pairs of medial setae.

Eyes stalked; antennule and antenna endopod first appear as buds; first two maxillipeds with 6 natatory setae on exopods; developing third maxiliiped may be present under carapace; no uropods; telson usually not separated from the 6th abdominal somite (e.g. except Majidae), furcal arch may bear 4 pairs of setae.

Eyes stalked; mandibular palp may be present as primordia; first two maxillipeds with 8 natatory setae on exopods; developing third maxiliiped usually present; developing pereopods and pleopods may be present; uropod buds present; telson separated from 6th abdominal somite; may have additional setae on furcal arch.

Eyes stalked; mandible with palp primordia; exopod of maxillipeds with 10 or more natatory setae; pleopod and pereopod buds more developed than previous stages but not yet functional; telson separated from 6th abdominal somite; may have additional setae on furcal arch.

Eyes stalked; mandible with setose palp; maxillipeds no longer natatory, now functional mouthparts; pereopods complete and functional, the first one chelate; pleopods biramous, segmented and setose; uropods setose.

Table 3. General characters to recognize the stages of brachyuran larval phases (exceptions apply, especially among lower Brachyura, e.g. Dromiidae, Homolidae and Raninidae).

The abdomen of the first zoeal stage is composed of five articulating somites and a terminal telson (by others the telson is considered a separate entity from the abdomen). Older zoeae acquire a sixth somite at some later stage, except in Leucosiidae and some Pinnotheridae. On abdominal somites 2 and 3 a pair of dorsolateral processes are present in almost all species. The posterolateral margins of somites 3-5 vary in shape from rounded to bearing subacute or acute processes. The pleopod buds arise ventrally on somites 2-6 in older zoeae.

The shape of the telson remains more or less constant throughout development. In the majority of crab larvae the posterolateral portions of the telson are extended into conspicuous furcae. The remaining posterior margin of the telson is formed into a slight median lobe in some Pinnotheridae, is convex in Leucosiidae but is variably concave in most other species. The posterior margin also bears plumodenticu-late setae. Zoeae of Dromiacea have ten setae, Dorippidae only two, and in all other brachyurans studied there are six setae in the first stages, with additional setae added in later stages during development

in the majority of brachyrhynchs. The outer lateral spine of the telson is present in nearly all previously studied species. The dorsal spine is absent in Dromiacea, Oxyrhynchs, but it is developed in all stages of most brachyurans.

The antennule is represented by a subcylindrical exopod with an undifferentiated protopod. The exopod bears aesthetascs and sometimes additional simple setae, whose number varies with the zoeal stage.

The antenna is well developed in zoeal stages of all species in the area, except for Zaops and Persephona, where it is represented only as a small bud. Three general types of antennae are distinguished". In Dromiacea occurs a broad, somewhat flattened, setose exopod, the endopod is subcylindrical and bears terminal setae; In Hymenosomatidae and some Grapsidae and Xanthidae species, the antennal exopod is absent or reduced to a small simple seta, small setose process or a minute articular process bearing 1-2 terminal simple setae;


In Parthenopidae, all other oxyrhynchs and brachy-rhynchs the antenna is more developed - the exopod is long and often extends to the distal half of the spinous process which is well developed and reaches from less than half the length to almost the tip of the rostrum. The endopod bud can be distinguished starting in the second stage.

The mandible is composed of distal incisor and molar processes and in some species an endopodite bud may occur basally in the last zoeal stage.

The maxillule is composed of an endopod and a pro-topod; from the latter arise the basis and coxa.

The maxilla is composed of four endites: endopod, basis and coxa, and the scaphognathite, which in all species is developed into an elongated lobe. The scaphognathite margin bears a consistent number of plumose setae in the first zoeal stage but the number of setae varies within a species in later stages.

The first maxilliped consists of a coxa that is often imperfectly demarcated from the adjoining basis which bears a characteristic setal pattern that in most species remains unchanged during zoeal development; from the distal portion of the basis arises an endopod and exopod. The setose endopod is 5-segmented in all species studied. The unsegmented, bisegmented or partially bisegmented exopod of all species has long plumose natatory setae distally, starting with four in the first stage, to as many as 13 in older zoeae (Brossi-Garcia and Rodrigues, 1993).

The second maxilliped is similar to the first except for the endopod which is 4-segmented in the early larval stages of Dromiacea and 3-segmented in other bra-chyuran zoeae studied. Exceptions include Zaops, where only two segments can be recognized, and in Persephona, in which the endopod is unsegmented.

The third maxilliped can appear under the carapace as a developing non-functional bifurcated and unsegmented structure in early zoeal stages of some species but it usually occurs in later stages in species with more zoeal stages.

Developing pereopods appear as gradually growing buds under the carapace during zoeal development.

The megalopa (Fig. 3)

The body of a megalopa is formed by an oval or rectangular cephalothorax, externally composed of a dorsal carapace and a ventral sternum, plus a segmented abdomen. Large stalked pigmented eyes and other appendages emanate from the cephalothorax, as do swimming appendages from the abdomen. Both the cephalothorax and abdomen are pigmented in live or freshly preserved specimens. This is not shown in the figures.

The carapace frequently has a rostrum, except in Zaops. The carapace can de ornamented by protuberances and bear a posteriorly directed median spine (Parthenope). The pereopods (pi, p2, p3, p4, p5) are fully formed, the first one being chelate. The abdomen is composed of six somites in all megalopae studied, except in Zaops and Persephona, in which only five somites are present, or the 6th is incipiently formed. Ventrally the abdomen bears functional biramous ple-opods on somites 2-5, and a pair of uniramous uro-pods on somite 6 in most species.

The antennule is developed as a stout 3-segmented peduncle bearing an endopod and exopod. The exopod segments are often armed with aesthetasc setae.

The antenna is composed of a peduncle (protopod) with 4 segments and a multisegmented flagellum (endopod).

The mandibles no longer have distinct molar and incisor portions but a cutting edge instead. There is a dorsal palp that usually bears setae.

The maxillule and maxilla have changed little since the zoeal stage, except that the endopod now is usually reduced.

The first maxilliped has a protopod from which arise broad coxal and basial segments. A narrow endopod and a 2-segmented exopod have developed. The outer margin of the protopod has an epipod with a variable number of plumodenticulate setae.

The second maxilliped has a small protopod from which arises an endopod with four clearly demarcated segments and a 2-segmented exopod. For known species there is no epipod emanating from the protopod.


CRAB ZOEAL MORPHOLOGY

/ dorsal spine

rostral spine

© Lateral view of a generalized zoea III

@ Appendages l

bud of third

maxilliped

second maxilliped first maxilliped

dorsolateral process

posterolateral process

dorsal spine r ou te r

lateral spine

/

furca

Dorsal view of abdomen

mandible

maxillule

antennule

antenna

maxilla


The third maxilliped has a protopod from which arises a well developed 5-segmented endopod and a narrow 2-segmented exopod. The protopod is usually broad, with the external margin bearing a well developed epi-pod covered with plumodenticulate and other setae.

The pereopods have seven segments in the majority of the brachyuran megalopae. However, the basial segment is not always clearly differentiated from the ischium. The lower margin of the dactyl on the 5th pereopod (p5) is armed with simple setae in almost all species studied.

The pleopods are attached to the ventral surface of the abdominal somites. Each appendage bears long marginal natatory plumose setae, the number of setae decreasing on succeeding pairs. A small endopod is present which is distally armed with a number of specialized setae that interlock with those of the adjacent pleopod and allow for synchronized movement of the swimming appendages.

General outline classification

The classification of brachyurans of the Southwest Atlantic Ocean is summarized as follows based on Bowman and Abele (1982), Williams (1984), and Melo (1996):

Superclass Crustacea Pennant, 1777 Class Malacostraca Latreille, 1806 Subclass Eumalacostraca Grobben, 1892

Superorder Eucarida Caiman, 1904 Order Decapoda Latreille, 1803 Suborder Pleocyemata Burkenroad, 1963 Infraorder Brachyura Latreille, 1803 Section Dromiacea De Haan, 1833

Superfamily Dromioidea De Haan, 1833 Family Dromiidae De Haan, 1833

Superfamily Homoloidea De Haan, 1839 Family Homolidae De Haan, 1839 Family Latreilliidae Stimpson, 1858

Section Oxystomata H. Milne Edwards, 1834 Superfamily Dorippoidea De Haan, 1833

Family Dorippidae De Haan, 1833 Superfamily Calappoidea De Haan, 1833

Family Calappidae De Haan, 1833 Family Leucosiidae Samouelle, 1819

Superfamily Cyclodorippoidea Ortmann, 1892 Family Cyclodorippidae Ortmann, 1892 Family Cymonomidae Bouvier, 1897

Superfamily Raninoidea De Haan, 1833 Family Raninidae De Haan, 1833

Section Oxyrhyncha Latreille, 1803 Superfamily Majoidea Samouelle, 1819

Family Majidae Samouelle, 1819 Superfamily Parthenopoidea MacLeay, 1838

Family Parthenopidae MacLeay, 1838 Superfamily Hymenosomatoidea MacLeay, 1838

Family Hymenosomatidae MacLeay, 1838 Section Brachyrhyncha Borradaile, 1907

Superfamily Cancroidea Latreille, 1803 Family Atelecyclidae Ortmann, 1893

Superfamily Bellioidea Dana, 1852 Family Belliidae Dana, 1852

Superfamily Portunoidea Rafinesque, 1815 Family Portunidae Rafinesque, 1815

Superfamily Xanthoidea MacLeay, 1838 Family Geryonidae Colosi, 1924 Family Xanthidae MacLeay, 1838 Family Goneplacidae MacLeay, 1838

Superfamily Pinnotheroidea De Haan, 1833 Family Pinnotheridae De Haan, 1833

Superfamily Grapsoidea MacLeay, 1838 Family Grapsidae MacLeay, 1838 Family Gecarcinidae MacLeay, 1838

Superfamily Ocypodoidea Rafinesque, 1815 Family Ocypodidae Rafinesque, 1815 Family Palicidae Bouvier, 1897

Superfamily Cryptochiroidea Paulson, 1875 Family Cryptochiridae Paulson, 1875

Identification

Table 4 summarizes the present knowledge of larval development for families of brachyuran crabs from the Southwest Atlantic Ocean. As mentioned above, it is important for the reader to realize that because of many unknown larvae, those to be identified may not fit any larvae described herein. However, in such cases the keys may help in narrowing the search to higher groups, such as families. It also must be recognized

"*—r Fig. 2. Crab zoeal morphology. Original.


CRAB MEGALOPAL MORPHOLOGY

cheliped

pereopod 2

pereopod 3

pereopod 4

pereopod 5

© Dorsal view of generalized megalopa

first maxilliped

maxilla

third maxilliped

( 3 Appendages second maxilliped


Section i

D ROM 1 ACE A

OXYSTOMATA

OXYRHYNCHA

BRACHYRHYNCHA

Family

Dromiidae

Homolidae

Latreilliidae

Dorippidae

Calappidae

Leucosiidae

Cyclodorippidae

Cymonomidae

Raninidae

Majidae

Parthenopidae

Hymenosomatidae

Atelecyclidae

Belllidae

Portunidae

Geryonidae

Xanthidae

Goneplacidae

Pinnotheridae

Grapsidae

Gecarcinidae

Ocypodidae

Palicidae

Cryptochiridae

TOTAL

Number

of species

4

2

2

4

12

20

8

4

5

83

16

1

1

4

24

2

53

18

18

24

2

12

7

2

328

Number

of genera

3

2

1

2

5

9

4

2

3

45 • • - J

8

1

1

3

9

1

26

13

9

15

2

3

1

2

170

Number of

species with

known larval

development

4

1

1

• 1 !

2

1 _

0

0

0

19

1

1

1

3

9

0

23 i

0 \

9

17

1

7

0

1

102

Table 4, Summary of brachyuran crabs from the Southwest Atlantic and the number of species with known larval development.

« -Kg. 3. Crab megalopal morphology. Original.


First zoeae of Dromiidae

0.5

Dromia erythropus (DV)

0.5

Cryptodromiopsis antillensis (DV)

Hypoconcha sabulosa (DV)

Cryptodromiopsis antillensis (A2,DV)

Fig. 4. First zoeae of Dromiidae. A2: antenna; DV: dorsal view; LV: lateral view; scale in mm. Sources, from Lang and Young (1980): C; from Laughlin et al. (1982): A; from Rice and Provenzano (1966): B-D.

First zoeae of Dromiidae

Species Carapace Abdomen Antennule

With blunt dorsal projection, Pair of blunt dorsal

Cryptodromiopsis antillensis transverse groove and two pairs of projections 2 aesthetascs

lateral grooves j on somites 2-5

Dromia erythropus No dorsal projection but with spine No dorsal projections

Hypoconcha sabulosa

Hypoconcha arcuata

on each posterolateral corner

No spine but one pair of transverse

grooves

No spines and no grooves

or spines

No spines

6 aesthetascs

Maxilliped 3

endopod

2 plumose

setae

1 denticulate

1 simple seta

3 aesthetascs 2 plumose

setae

No spines 5 aesthetascs no setae

Table 5. Species characters of first zoeae of Dromiidae.

that most of the larval accounts are based on laboratory rearings, and it is still unclear how much variability there is between specimens obtained from the wild and those obtained from culture (Ingle, 1992). Thus definitive identifications should be obtained by consultations with experts in the field.

In order to arrive at an identification, the reader should first use the family key. Larvae are grouped using a minimum of obvious characters to separate families, so that in almost all cases a dissecting microscope is not necessary for rapid identification. Characters given in the appropriate family tables (Tables 5-13) can then


be used to further identify a specimen to the species level. Once a larva has been identified, we suggest that this be verified using additional details given in the original literature cited in Table 14.

It should be noted that, for practical purposes, the ordering of families does not exactly follow established groupings of adults.

Identification of crab zoeae: Key to the brachyu-ran families from the Southwest Atlantic based on the first larval stage

1 Carapace long, horizontally elongated or shrimp-like, telson subtriangular 2

la Carapace not long or shrimp-like, telson not sub-triangular (except Leucosiidae) 4

2 Carapace without dorsal and lateral spines; posterior margin not denticulate; antennal spinous process (protopod) shorter than endopodite: Dromiidae (Fig. 4, Table 5)

2a Carapace with dorsal and lateral spines; posterior carapace margin denticulate; antennal protopod longer than endopodite 3

3 Antennal spinous process longer than exopod; abdominal somites with dorsal and lateral spines, telson with dorsal spines: Homolidae (Fig- 5)

3a Antennal spinous process shorter than exopod; abdomen lacking dorsal and lateral spines and telson without dorsal spines: Latreilliidae (Fig. 6)

4 Carapace spines extremely long, more than twice carapace length; telson very long, exceeding length of abdominal somites, bearing two medial setae on furcal arch and pair of setae laterally on proximal portion of telson: Dorippidae (Fig. 7)

4a Carapace spines and telson shorter; telson seta-tion different 5

5 Telson sub-triangular, with short furcal spines and lacking medial arch; antenna rudimental, consisting of single process: Leucosiidae (Fig. 8)

5a Telson not sub-triangular, with longer distinct furcal spines and discrete medial arch; antenna more developed, usually consisting of more numerous and/or longer armed processes 6

6 Carapace with rostral, dorsal or lateral spines and conspicuous projection: Majidae (in part, Fig. 9, Table 6)

First zoea of Homolidae Homola barbata

(A2, DV)

Fig. 5. First zoea of Homolidae. A2: antenna; DV: dorsal view; LV: lateral view; scale in mm. Sources, from Rice and Provenzano (1970): A, B.

6a Carapace usually with the full complement of dorsal, rostral and lateral carapace spines, but without other conspicuous projection 12

6b Carapace lacking some spines 7 7 Carapace with all spines missing: Pinnotheridae

(in part - Z. ostreum, Fig. 10; Table 7) 7a Carapace with dorsal and/or lateral spines missing


9a 10

10a 11

11a

12

12a

13

13a

14

14a

15

15a

Carapace with rostral spine but lacking lateral and dorsal spines; antenna reduced to a protopod process with short proximal seta representing exopod; telson furcae shorter than telson body: Hymenosomatidae (Fig. 11) Carapace with rostral and dorsal spines but lacking lateral spines; antenna usually more developed and telson furcae longer 9 Rostral and dorsal carapace spines very long, at least 1.5x carapace length: Belliidae (Fig. 12, Table 8) Rostral and dorsal carapace spines shorter 10 Rostral spine shorter than antenna: Majidae (in part, Fig. 9, Table 6) Rostral spine longer than antenna 11 Antennal exopodite usually not well developed, less than Vi protopodite length: Grapsidae (Fig. 13, Table 9) Antennal exopodite usually well developed, more than Vi protopodite length: Ocypodidae (in part, Fig. 14, Table 10) Abdomen laterally and/or posteriorly expanded on some somites 13 Abdomen usually with more or less parallel sides and posterolateral spines shorter than somite 16 Abdominal somites 3-5 with prominent posterolateral spines, about as long as length of somite; antennal exopod shorter than protopod; telson furcae arched: Parthenopidae (Fig. 15) Abdominal somites with shorter posterolateral spines; antennal exopod either as long as spinous process or reduced to a seta; telson furcae parallel 14 Antennal exopodite reduced to a seta; abdominal somite 5 laterally expanded into wing-like extension: Pinnotheridae (in part- Pinnixa spp., Fig. 10, Table 7) Antennal exopodite not reduced to a seta, at least Vi length of spinous process; abdominal somite 5 not laterally expanded into wing-like extension

15 Abdominal somite 4 laterally expanded into winglike extension; antennal exopod about as long as spinous process: Cryptochiridae (Fig. 16) Abdominal somites 4 posterolaterally slightly expanded (more so and with rough surface in alter stages); antennal exopod less than V2 length of spinous process: Ocypodidae (in part, Ocypode quadrata, Fig. 14)

First zoea of Latreilliidae Latreillia elegans

Fig. 6. First zoea of Latreilliidae. A2: antenna; DV: dorsal view; LV: lateral view; scale in mm. Sources, from Rice and Williamson (1977): A, B (originally fig. 11B, C, Don p. 36).

16 Telson furcae with distinct spines 18 16a Telson without distinct spines 17 17 Antenna with protopod and well developed exo

pod process, latter about 3A length of spinous process: Gecarcinidae (Fig. 17)

17a Antenna with exopod shorter: Grapsidae (in part, Fig. 13)


17b Antenna without exopod: Pinnotheridae (in part, Fig. 10, Table 7)

18 Telson usually with two or three spines on each furca 19

18a Telson usually with single spine on each furca: Xanthidae (Fig. 18, Table 11)

19 Antennal exopod more than V2 length of proto-pod: Atelecyclidae (Fig. 19)

19a Antennal exopod equal or less than XA length of protopod: Calappidae and Portunidae (Fig. 20 and 21, Tables 12 and 13)

First zoea of Dorippidae Ethusa microphthalma

( I DV)

Fig. 7. First zoea of Dorippidae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Martin and Truesdale (1989): A-C.

First zoea of Leucosiidae Persephona mediterranea

© (A2, DV)

0.1

0.2

(T DV)

Fig. 8. First zoea of Leucosiidae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Negreiros-Fransozo etal (1989): A-C.


First zoeae of Majidae

Libidoclaea granaria (LV)

Taliepus dentatus (LV)

Anasimus latus (LV)



Stenorhynchus seticornis (T, DV)

Fig. 9. First zoeae of Majidae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Bakker et al. (1990): B; from Bolanos et al. (1996): F; from Campodonico and Guzman (1972): E; from Fagetti (1969): G; from Fagetti and Campodomco (1971): H; from Fransozo and Hebling (1982): D; from Fransozo and Negreiros-Fransozo (1997): L; from Hartnoll (1964): J; from Hiyodo et al. (1994): M; from Negreiros-Fransozo and Fransozo (1991): A; from Sandifer andvanEngel (1972): I; from Wilson et al. (1979): C; from Yang (1976); K,N, O.


S ubf ami I ies/S pecies


Spines and Dorsolateral

posterolateral projections

setae on carapace | on abdomen

Posterolateral ! Marginal + apical

processes j setae on

on abdomen [ scaphognathite

INACHINAE

Eurypodius latreillei

Stenorhynchus seticornis

INACHOIDINAE

Anasimus latus

Pyromaia tuberculata

TYCHINAE

Pitho Iherminieri

EPIALTINAE

Taliepus dentatus

Epialtus brasiliensis

Acanthonyx scutiformis

Epialtus bituberculatus

PISINAE

Libinia spinosa

Libidoclaea granaria

Libinia ferreirae

MITHRACINAE

Mithraculus forceps

Mithraculus coryphe

Mithrax verrucosus

Mithrax caribbaeus

Mithrax hispidus

Microphrys bicornutus

1 rostral, 1 dorsal,

2 laterals, 4 setae

1 dorsal, 3(4) setae

1 dorsal, several setae

1 dorsal

1 rostral, 1 dorsal, 7 setae



4 (5) setae

1 rostral, 1 dorsal


1 rostral, 1 dorsal



6(7) setae



! 1 rostral, 1 dorsal, 3 setae




2nd

2nd

2nd

and 3rd somites

and 3rd somites

2nd somite

2nd somite

and 3rd somites

2nd somite

2nd somite

2nd somite

2nd somite

2nd somite

2nd somite

2nd somite

2nd somite

2nd somite

2nd somite

2nd somite

2nd somite

2nd somite

3rd and 4th somites

3rd and 4th somites

3rd and 4th somites

absent

3rd and 4th somites

absent

absent

absent

absent

, 3rd and 4th somites

3rd and 4th somites

I 3rd and 4th somites

3rd and 4th somites

3rd and 4th somites

3rd and 4th somites

3rd and 4th somites

3rd and 4th somites

3rd and 4th somites

10

10 + 1

10 + 1

10 + 1

26

10 + 1

12 + 1

11 + 1

12 + 1

10

14 + 1

10+1

13

12 + 1

12 + 1

12 + 1 i

13

12(13) + !

Table 6. Species characters of first zoeae of Majidae.


First zoeae of Pinnotheridae

Fig. 10. First zoeae of Pinnotheridae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Costlow and Bookhout (1966b): F; from Dowds (1980): C, D; from Hyman (1925): A, B; from Marques and PoHe (1996b): G, H; from Pohle and Telford (1981): E, I.


Species

Clypeasterophilus stebbingi

Dissodactylus crinitichelis

Pinnaxodes chilensis

Pinnixa chaetopterana

j Pinnixa cristata

Pinnixa patagoniensis

Pinnixa sayana

j Tumidotheres maculatus

Carapace

all spines present;

knobs absent; D<1.5 x R

all spines present;


all spines present-

knobs absent; D^1.5 x R

all spines present-


all spines present;

knobs absent; D<1.5 x R;

all spines present;

knobs absent; D<1.5 x R;

all spines present;


all spines present;

knobs present; D<1.5 x R

Dorso

lateral

knobs

Telson

only on 2nd furcae spinulose

2nd and 3rd

2nd and 3rd

2nd and 3rd

2nd and 3rd

2nd and 3rd

furcae spinulose

furcae spinulose

median notch

median notch

median notch

2nd and 3rd furcae spinulose

2nd and 3rd furcae spinulose

Antenna

l imited to spinous process

limited to spinous process



+ 1 seta


+ 1 seta

• limited to spinous process

+ 1 seta

limited to spinous process + 1 seta


Zaops ostreum spines absent; knobs absent no spines \ median notch rudimentary

Table 7. Species characters of first zoeal stage Pinnotheridae (D: dorsal spine; R: rostral spine; CL: carapace length).

©

First zoea of Hymenosomatidae Halicarcinus planatus

0.5

(A2, LV)

0.2

0.2

(T, DV)

Fig. 11. First zoea of Hymenosomatidae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Boschi et al.

(1969): A-C.


First zoeae of Belliidae

(A2, LV)

Fig. 12. First zoeae of Belliidae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Boschi and Scelzo (1970): A-C; from Fagetti and Campodonico (1970): D-F.

Species

Acanthocyclus albatrossis

Acanthocyclus gayi

Corystoides chilensis

First zoeae of Belliidae

Carapace

with spinules only

on rostral and dorsal spines

with spinules on

spines and all over carapace

with spinules only on

rostral and dorsal spines

posterodorsal border

of 2nd - 5th

abominal somite

with minute spines + 2 setae

with minute spines + 2 setae

2 setae only

Table 8. Species characters of first zoeae of Belliidae.


First zoeae of Grapsidae

Percnon gibbesi (LV)

0.5

Goniopsis cruentata (LV)

0.5

Armases angustipes (LV)

0.5

Plagusia ,.„, depressa (LV)

Cyclograpsus integer (LV)

Pachygrapsus Y\ Eucniro„rapsus

transversus (LV) amencanus (LV) Cyrtograpsus altimanus (LV)

Planes cyaneus (LV) Crapsus

grapsus (LV)

yvlefasesarma rubripes (LV)

Chasmagnathus granulata (LV) Sesarma

rectum (LV)


Zoeae of Grapsidae

Subfamilies/Species

GRAPSINAE

Goniopsis cruentata

Grapsus grapsus

Pachygrapsus gracilis \ i

Pachygrapsus transversus

Planes cyaneus \

Geograpsus lividus

SESARMINAE

Aratus pisonii

Chasmagnathus granulata

Cyciograpsus integer

Metasesarma rubripes

Sesarma rectum

Armases (= Sesarma) angustipes

VARUNINAE

Cyrtograpsus altimanus

Euchirograpsus americanus

Cyrtograpsus anguiatus

PLAGUSIINAE

Plagusia depressa

Percnon gibbesi

Antennal endopod

length in relation

to protopod

rudimentary

half !

rudimentary

rudimentary

rudimentary

rudimentary

half

half

equal

half

half

half

half

rudimentary

half

rudimentary

rudimentary

Carapace

lateral spines

knob-like

absent

absent

knob-like

absent

absent

absent

present

absent

absent

absent

absent

present

present

present

present

present

Maxilla

endopod1

setation

2,2

2,3 ;

2,2 :

2,2

2,2

2 ,2

2,3

2,2

2,2

2,3

2,3

2,3

2,2

: 2 ,3

2,2

2,2 r

i..... 2.ijL

Maxilliped 1 i

basipod

setation

2, 2, 2, 2

1

2, 2, 2, 2

2 , 2 , 2 , 2

2, 2, 2, 2

2, 2, 2, 2

2, 2, 3, 3

2, 2, 2, 2

2 , 2 , 3 , 2

2, 2, 3, 3

2 , 2 , 3 , 3

2, 2, 3, 3

2, 2, 3, 3

2, 2, 2, 2

2, 2, 3, 3

2 , 2 , 2 , 2

2, 2, 3, 2

Maxilliped 1

endopod

setation

1 , 2 , 1 , 2 , 5

1

1 , 2 , 1 , 2 , 5

1 , 2 , 1 , 2 , 5

1,2, 1,2,5

1 , 2 , 1 , 2 , 5

2 , 2 , 1 , 2 , 5

1 , 2 , 1 , 2 , 5

2 , 2 , 1 , 2 , 5

2 , 2 , 1 , 2 , 5

2, 2, 1,2, 5

2 , 2 , 1 , 2 , 5

2 , 2 , 1 , 2 , 5

2 , 2 , 1 , 2 , 5

2 , 2 , 1 , 2 , 5

2 , 2 , 1 , 2 , 5

A 2 ' ! ' 2 / !

Table 9. Species characters of zoeae of Grapsidae.

Fig. 13. First zoeae of Grapsidae. LV: lateral view; scale in mm. Sources, from Boschi et al. (1967): M, 0 (part); from Cuesta and Rodriguez (1994): F; from Diaz and Ewald (1968): L (originally fig. 1 on p. 226); from Fransozo and Hebling (1986): N; from Fransozo et al. (in press): E; from Gore and Scotto (1982): D, O (part); from Gore et al. (1982): O (part); from Konishi and Minagawa (1990): J; from Kowalczuk (1994): I; from Lewis (1960): K; from Paula and Hartnoll (1989): A, O (part); from Warner (1968): B; from Wilson (1980): G, O (part); from Wilson and Gore (1980): C.


First zoeae of Ocypodidae

Ocypode quaarata (LV)

Ucides cordatus (LV)

©

(A2, DV) (T, DV)

Uca uruguayensis

Fig. 14. First zoeae of Ocypodidae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Diaz and Costlow (1972): A; from Rieger (1996): B, D-E; from Rodrigues and Hebling (1989): C.

Zoeae of Ocypodidae

Uca species

U. uruguayensis

U. vocator

U. mordax

U. burgersi

U. thayeri

Genera

Ocypode

Ucides

Lateral spine on carapace: Antennal exopod setation

Uca j

Antennule setation

present

absent

absent

Maxillule

2(3) aesthetascs + 1(2) setae

2 aesthetascs + 1 (2) setae

2(3) aesthetascs + 1(2) setae

2(3) aesthetascs +

2 aesthetascs +

1 seta

seta

l

r

protopod

1 seta

no seta

no seta

no seta

no seta

2 setae

2 setae

3 setae

Maxilliped 1 setation;

endopod setation

2 , 2 , 1 , 2 , 5

2 , 2 , 1 , 2 , 5

| 2 , 2 , 1 , 2 , 5

2 , 2 , 1 , 2 , 5

0 , 1 , 1 , 1 , 4 1

Number of abdominal

dorsal setae

1 , 2 , 2 , 2 , 2

0 , 2 , 2 , 2 , 2

0, 2, 2, 2, 2

0, 2, 2, 2, 2

0, 2, 2, 2, 2 I

Table 10. Generic and species characters of zoeae of Ocypodidae.


First zoea of Parthenopidae First zoea of Cryptochiridae Parthenope (Platylambrus) serrata Troglocarcinus coral I i cola

Fig. 16. First zoea of Cryptochiridae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Scotto and Gore (1981): A-C.


First zoea of Gecarcinidae Cardisoma guanhumi

(A2, DV) (T, DV)

Fig. 17. First zoea of Gecarcinidae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Costlow and Bookhout (1968): A-C.


First zoeae of Xanthidae

Hexapanopeus schmitti (LV)

(A2, LV)

Panopeus bermudensis

Group C

0.2

(LV) \ \ (A2,DV)

Micropanope scuiptipes

Group B

0.5

Eurytium limosum (LV)

Eurypanopeus abbreviates (LV)

Xanthodius denticulatus (LV) Garthiope barbadensis (LV)

(A2, DV) Hexapanopeus paulensis

0.5

Panopeus americanus (LV)


First zoeae of Xanthidae Group D

© ^ ®

Eriphia gonagra (LV) Carpilius corallinus (l_V) Xanthodius parvulus (LV)

Platyxanthus crenuiatus (LV)

Menippe nodifrons (LV)

0.5

(A2, DV)

Pilumnus dasypodus

Fig. 18. Zoeae of Xanthidae. A2: antenna; DV: dorsal view; LV: lateral view; scale in mm. Sources, from Andryszak and Gore (1981): I, L; from Bakker et al. (1989): A; from Bookhout and Costlow (1979): S; from Fransozo (1987): N; from Fransozo et al. (1989): F, H; from Gore et al. (1981): K; from Kurata et al. (1981): D; from Laughlin et al. (1983): O; from Lebour (1944): J, P; from Martin et al. (1985): B, C; from Menu-Marque (1970): M; from Negreiros-Fransozo (1986a): E; from Negreiros-Fransozo (1986b): G; from Sandifer (1974): R; from Scotto (1979): Q.


Zoeae of Xanthidae

SPECIES

Panopeus

herbstii

Panopeus

bermudensis

Medio-dorsal Lateral

Sharp | -E abdominal j •% + a. co l.f,

" • 1 : W

Telson Maxi l l l iped 1 : Maxi l l l i - furcae:

basis + : ped 2 Telson dorsal, knob on spines on P™?*™' j | | l i ^ l ^ l f l i endopod basis + furcal outer

I .= ! ? S * = .2 setation

lateral process

carapace carapace ^ ^ ^ = i H j j ' S | = " ' -g = setation endopod surface and !

i-H I H 111

inner spines

absent

absent

present ; somites 3-5; 3 + 2 1,6 4 , ?;3, 2, 1,2, 5 ? ; 1 , 1 , 5 smooth 1, 2, 0

absent absent ! 3(4) + 0; a | 1, 6 4

Panopeus

occidentalis

|Panopeus

j americanus

I Panopeus

present

absent

present in

9; 3, 2, 1 ,2, 4 3; 1 , 1 , 3 : smooth 1 , 0 , 0

10; 3, 2, 1, 2, 5 4 ; 1, 1, 5 j spinulose 1, 2, 0

10; 3, 2, 1,2, 5 4 ; 1 , 1 , 5 1 smooth 1 1 , 1 , 0

absent

absent

present only

in zoea 3-4

austrobesus j later zoeae

Hexapanopeus

paulensis _

Hexapanopeus

angustifrons

! Hexapanopeus

\ caribbaeus

Hexapanopeus

• schmitti

Eurypanopeus

i abbreviatus

' Eurypanopeus

depressus

Pilumnus

dasypodus

Eriphia

gonagra

Menippe

nodifrons

Eurytium

limosum

; Carpilius

• corallinus

• Cataleptodius

floridanus

present j somites 3-5 3 + 1 ' b I 1,6 3

J _ J _ _±_ ! i " ' i " 'j " : "

present (somites 3-5 3 + 1 i b I 1,6 4 _ I L I I i : : :

present somites 3-5 3 + 1 I b [ 1 , 6 4 10; 2, 2, 1, 2, 5 4; 1 , 1 , 4 smooth 1 , 2 , 0

1,6 4 10; 3, 2, 1,2,5J 4; 1 ,1 ,5 smooth j 0, 0, 0

1,6 4 4?; 3, 2, 1, 2, 5 3?; 1, 1, 5 smooth 0, 0, 0

1,6 4

present absent 3 + 1

present present absent

present somite 5

present as a absent , absent

protuberance

present somites 3-5

4 + 1

2 + 2

2 + 1

2 + 1

only zoea 4 present l somites 3-5

present

absent

present

1,5 4

10; 3, 2, 1 ,2, 5! 4 ; 1 , 1, 5 j smooth 1 , 0 , 0 j_ J

10; 3, 2, 1, 2, 5 2; 1 , 0, 4 . smooth , 0, 1, 1

1,6 \ 4

2 + 1 1,6 5

present

absent ; 2 + 2 ' c i 1, 6 • 4

absent

9; 3, 2, 1 ,2, 5 4 ; 1 , 1 , 4 j smooth 1 , 2 , 0

8; 3, 2, 1 , 2, 5 , 4 ; 1 , 1, 5 smooth ; 1, 0, 0

10; 3, 2, 1 ,2 ,5 4; 1, 1, 6 ' spinulose1 1 , 2 , 0

3 + 0

present present ] somite 5 3 + 1

c 1,6 4 i 1 0 ; 2 , 2, 1,2, 5| 4; 1, 1, 6 : s m o o t h : 1 , 2 , 0

d 1,4 4

present

present

absent

present absent 2 + 3 b I 1, 6 ?

10; 3, 2, 1,2, 5; 4 ; 0 , 1,4 j smooth j 1,2, 0

?;3, 2, 1,2, 5 ?;? smooth 1 , 2 , 0 i

present | absent ; 2 +1

present absent

Garthiope

: barbadensis

Micropanope

sculptipes

: Platyxanthus

crenulatus

Platyxanthus

patagonicus

Xanthodius

denticulatus

3 + 1

1,6 4 8; 2, 2, 1,2, 5 ?; 1 , 1 , 4 ; ? j 0, 2, 0

present present i absent ! 2 + 2

1,5 4 j 10; 3, 2, 1, 2, 5 4; 1 , 1 , 5 spinulose 1, 2, 0

1 , 6 | 4 1 0 ; 2 , 2 , 1,2, 5 4 ; 1, 1, 5 smooth 1 , 2 , 0

present

absent

present

absent

present

present

present

present

Xanthodius

parvulus

somites 3-51 3 + 2

absent ; 2 + 2

somites 3-5; 2 + 2

somites 3-5! ?

1,6 4 M 0 ; 2, 2, 1,2, 5! 4; 1, 1,5 smooth j 1 , 1 , 0

1,6 4 8; 3, 2, 2, 1, 4 4; 1, 1, 5 smooth

1 , 6 : 4 10; 3, 2, 1,2, 5 : 4 ; 1, 1,6 smooth

1,0, 0

, 2 , 0

absent present j absent ?

?;? i smooth 1, 2, 0

?; ? ! smooth 1, 2, 0

Table 11. Species characters of zoeae of Xanthidae.


First zoea of Atelecyclidae Peltarion spinulosum

0.2

(T, DV)

(A2, LV)

Fig. 19. First zoea of Atelecyclidae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, ftom lorio (1983): A-C.


First zoeae of Calappidae

(A2, DV)

Ca lappa gal I us

0.2

Fig. 20. First zoeae of Calappidae.A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Rieger and Hebling (1993): D-F; from Taishaku and Konishi (1995): A-C.

First zoeae of Calappidae

Species Rostral spine

Hepatus pudibundus smooth

Calappa gallus 15-10 spinules on anterior surface

Table 12. Species characters of first zoeae of Calappidae.


First zoeae of Portunidae

Fig. 21. First zoeae of Portunidae. A2: antenna; DV: dorsal view; LV: lateral view; T: telson; scale in mm. Sources, from Bookhout and Costlow (1974): E, H; from Boschi (1981): A; from Costlow and Bookhout (1959): D; from Negreiros-Fransozo (1996): G; from Stuck and Truesdale (1988): C; from Wear and Fielder (1985): B (originally fig. 130 on p. 51), F (orig. fig. 134 on p. 53), I (orig. fig. 127 on p. 51).


First zoeae of Portunidae

Subfamilies/Species Abdominal somites 3-5 Telsonfurcal spines i

PORTUNINAE

Arenaeus cribrarius I serrate lateral projections

Callinectes sapidus

Portunus spinicarpus

prominent lateral spines

overlaping the adjacent somite



Scylla serrata short lateral spines

Charybdis hellerii

POLYBIINAE

Coenophthalmus tridentatus



r short lateral spines

Ovalipes trimaculatus

Ovalipes catharus

short lateral spines

Ovalipes punctatus



Antennule

setation

Maxilliped 1

endopod

setation

1 lateral, 1 dorsal j

1 lateral, 1 dorsal j

1 lateral, 1 dorsal

1 lateral

3 aesthetascs +

| 3 simple setae

3 aesthetascs +

I 2 simple setae

3 aesthetascs +

3 simple setae

3 aesthetascs +

1 simple seta

2, 2, 0, 2, 5

2, 2, 0, 2, 5

2 , 2 , 0 , 2 , 5

2, 2, 0, 2, 2

1 lateral, 1 dorsal

1 dorsal

2 aesthetascs +|

1 simple seta 3(2), 2, 0, 2(3), 5(6)

1 lateral, 1 dorsal

1 lateral, 1 dorsal

1 lateral, 1 dorsal

3 aesthetascs +

1 simple seta

2 aethetascs +

3 simple setae 2 , 2 , 1 , 2 , 5

Table 13. Species characters of first zoeae of Portunidae.


Species

FAMILY DROMIIDAE

Cryptodromiopsis antillensis

Dromia erythropus

Hypoconcha arcuata

Hypoconcha sabuiosa

FAMILY HOMOLIDAE

Homola barbata

FAMILY LATREILLIIDAE

Latreillia elegans

FAMILY DOR1PPIDAE

Ethusa micropbthalma

FAMILY CALAPPIDAE

Calappa gallus

Hepatus pudibundus

FAMILY LEUCOSIIDAE

Persephona mediterranea

FAMILY MAJIDAE

Acanthonyx scutiformis (as A. petiverii

; Anasimus latus





Libinia ferreirae

Libinia spinosa

Macrocoeloma camptocerum

Microphrys bicornutus

Mithraculus coryphe

| Mithraculus forceps

Mithrax caribbaeus

Mithrax hispidus

Mithrax verrucosus *

Pitho Iherminieri

• Pyromaia tuberculata


Taliepus dentatus

Num

larva! 1

Zoea \

6(7)

5

3 !

3

7

2

4

• _

1 +

4 (5) (6) |

4

I 2 j I

i 2 i i

2 ;

2

2

; 2 !

2 ;

2 I i

2 ;

2

2

2

2

2

2

2

2

2

2

ber of

stages

Megalopa

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

Described by

Rice and Provenzano, 1966

jLaughlin et al., 1982

Kircher, 1970

Lang and Young, 1980

Rice and Provenzano, 1970

Rice and Will iamson, 1977; Rice,

Martin and Truesdale, 1989

Taishaku and Konishi, 1995

| Rieger and Hebling, 1993

i Negreiros-Fransozo et al., 1989

i Hiyodo et ai., 1994

iSandifer and van Engel, 1972

Negreiros-Fransozo and Fransozo,

Negreiros-Fransozo and Fransozo,

Campodonico and Guzman, 1972

Fagetti, 1969

Bakkeretal. , 1990

Boschi and Scelzo, 1968

Yang, 1967

Hartnoll, 1964; Gore et al., 1982

Scotto and Gore, 1980

Wilson eta l . , 1979

Bolanos et al., 1990

Fransozo and Hebling, 1982

Bolanos and Scelzo, 1981

Bolanos eta l . , 1996

Fransozo and Negreiros-Fransozo,

Yang, 1976

: Fagetti and Campodonico, 1971

1982

1996

1991

1997

- —

.... _ j

-- -

—


Species

Number of

larval stages

Zoea Megalopa

Described by

FAMILY PARTHENOPIDAE

Parthenope (Platylambrus) serrata

FAMILY HYMENOSOMATIDAE

Halicarcinus planatus

Yang, 1971

1 Boschi etal . , 1969

FAMILY ATELECYCLIDAE

Peltarion spinosulum

FAMILY BELLIIDAE

Acanthocyclus albatrossis

Acanthocyclus gayi


FAMILY PORTUNIDAE

Arenaeus cribrarius

Callinectes sapidus

Charybdis hellerii

Coenophthalmus tridentatus

Ovalipes catharus

Oval ipes punctatus

Ovalipes trimaculatus


Scylla serrata

FAMILY XANTH IDAE

Carpilius corallinus

Cataleptodius floridanus

Eriphia gonagra

Eurypanopeus abbreviatus

Eurytium limosum

Euypanopeus depressus

I Garthiope barbadensis

Hexapanopeus angustifrons

Hexapanopeus caribbaeus

Hexapanopeus paulensis

Hexapanopeus schmitti

Menippe nodifrons

Micropanope sculptipes

Panopeus americanus

Panopeus austrobesus

4

4

4

4

8

7(8)

1 +

1 +

8

6

1 +

7

5

5

1

4

4

4

4

3(4) I

4 :

4

4

4

5(6)

4

4

4

1

1

1

1

1

1

1 *

1*

1

1

i *

1

1

1* i *

1

1

1

1

1

1

1

1

1

1

1

1

1

lorio, 1983

Campodonico and Guzman, 1973

Fagetti and Campodonico, 1970

Boschi and Scelzo, 1970

Stuck andTruesdale, 1988

Costlow and Bookhout, 1959

Negreiros-Fransozo, 1996 i

Boschi, 1981

Wear and Fielder, 1985

Terada, 1980

Boschi, 1981

Bookhout and Costlow, 1974

Ong, 1964; Wear and Fielder, 1985

Laughlin eta l . , 1983

Ingle, 1987 \

i Fransozo, 1 987

Negreiros-Fransozo, 1986a

Kurata et a!., 1981; Messerknecht et al., 1991

Costlow and Bookhout, 1961 a

Gore eta l . , 1981

Costlow and Bookhout, 1966a

Vieira and Rieger, in press

Fransozo et al., 1989

Bakker eta l . , 1989

Scotto, 1979

Andryszak and Gore, 1981

Negreiros-Fransozo, 1986b

Montu eta l . , 1988


Species

....

Panopeus bermudensis

Panopeus herbstii

Panopeus occidentalis

Pilumnus dasypodus

Platyxanthus crenulatus

Platyxanthus patagonicus

Xanthodius denticulatus

Xanthodius parvulus

FAMILY PINNOTHERIDAE


Dissodactylus crinitichelis

Pinnaxodes chilensis

Pinnixa chaetopterana

Pinnixa cristata

Pinnixa patagoniensis

Pinnixa sayana

Tumidotheres maculatus

Zaops ostreum

FAMILY GRAPSIDAE

Aratus pisonii

Armases angustipes

Chasmagnathus granulata

Cyclograpsus integer

Cyrtograpsus angulatus

Cyrtograpsus altimanus

Euchirograpsus americanus

Goniopsis cruentata

Crapsus grapsus = G. adcensionis

] Metasesarma rubripes

Pachygrapsus gracilis

Pachygrapsus transversus _

1 Percnon gibbesi Plagusia depressa

Planes cyaneus

Planes minutus

| Sesarma rectum

Number of

larval stages

Zoea Megalopa

4 ' 1

4

4 1

4

4 1

4 1

1+ i 1 *

1+ | 1 *

4

3 '

" 1+ 1*

1 +

1+ ! ?

5 1

1+ ! ?

| 5 1

I 4 j 1 !

4 | 1

! 4 1

; 4(5) ! 1

5(6) ! 1

5 1

5 1

5(6) 1

1+ i 1*

1 +

5

13

11

6

1 *

1 *

5 ?

! 1+ ! 1 *

2 1

3

Described by

Martin eta l . , 1985

Costlow and Bookhout, 1961b

Ingie, 1985

Sandifer, 1974; Bookhout and Costlow, 1979

Menu-Marque, 1970; Boschi, 1981

lorio and Boschi, 1986

Lebour, 1944

Lebour, 1 944

Marques and Pohle, 1996b

Pohle and Telford, 1981

Gutierrez-Martinez, 1971

Dowds, 1980; Sandifer, 1972

Dowds, 1980

Boschi, 1981

Dowds, 1980

Costlow and Bookhout, 1966b

Hyman, 1925; Sandifer, 1972;

Sandoz and Hopkins, 1947

Warner, 1968; Diaz and Bevilacqua, 1987

Kowalczuk, 1 994

Boschi e ta l . , 1967

Gore and Scotto, 1982

Rieger and Vieira, 1997

Scelzo and de Bastida, 1979

Wilson, 1980

Fransozo eta l . , 1998

Lewis, 1960

Diaz & Ewald, 1968; Montu et al., 1990

Ingle, 1987; Brossi-Ga"rcia and Rodrigues, 1993

;Cuesta and Rodrigues, 1994

: Paula and Hartnoll, 1989

Wilson and Gore, 1980

Konishi and Minagawa, 1990

Lebour, 1944

Fransozo and Hebling, 1986


Species

FAMILY GECARCINIDAE

Cardisoma guanhumi

FAMILY OCYPODIDAE

Ocypode quadrata

Uca burgers!

Uca mordax

Uca thayeri

Uca uruguayensis

Uca vocator

Ucides cordatus

FAMILY CRYPTOCHIRIDAE

Troglocarcinus coraliicoia

Number of

larval stages

Zoea

5

i

5

5(6) !

- 5(6) j

-M 5(6)

4(5-6) 5(6)

1

5

Megalopa

1

1

1

1

1

1

1

1

?

Described by

Costlow and Bookhout, 1968

Dfaz and Costlow, 1972

Rieger, 1998

; Rieger; 1992 ! Anger etal . , 1990

i Rieger, 1996

! Rieger, 1992

; Rodrigues and Hebling, 1989 i

Scotto and Gore, 1981

Table 14. Southwest Atlantic species of brachyuran crabs with known larval development and respective references. +: with additional unde-scribed stages; *: incomplete description; ?: unknown. Numbers in brackets include optional additional stage(s).

Identification of crab species in the megalopa phase

This last larval stage is the least well known among crab larvae, unrecognized in some species with known zoeal stages and being poorly described in other cases. While megaiopae show many distinct features, these characters are often not suitable for identification because they are not available for enough species to be useful for comparative purposes. There are also few obvious features that readily characterize members of specific families. This complicates identification to a point that does not allow for the construction of keys that are easy to follow. Thus, for identification purposes the reader should instead note Ihe size and gross morphological features of collected specimens (see below), and then compare these with the illustrations provided (Fig. 22-27). Subsequently, Tables 15-19 can be used for identification verification, by checking patterns of appendage setation.

Gross-morphological features that should be used in the identification process include the following:

1. Relative size of specimens in terms of carapace length or width, some being much larger (e.g.

2.

3.

Ocypode quadrata) than others (e.g. Clypeastero-philus stebbingi). Relative shape of the carapace, by comparing length to width, and the shape and width of the anterior margin between eyes (e.g. contrast Aratus pisonii, Armases angustipes, and Acanthonyx scutiformis). Ornamentation of the carapace; this can range from a smooth surface (e.g., Dissodactylus crinitichelis), hairy appearance (Cryptodromiopsis antillensis), to cuticular projections in specific locations that range from rounded knobs (Mithrax hispidus) to "antlers" (Latreillia elegans) and spines of various lengths (e.g., compare Libinia spinosa and Tumidotheres maculatus). Characteristics of pereopods: note the relative length of walking legs to the carapace (e.g., very long for Stenorhynchus, short in Libinia and Taliepus), the absence or presence of long swimming setae on the dactyls of the last pair of legs (e.g., compare Dissodactylus crinitichelis and Tumidotheres maculatus), and if the same dactyl is turned upward (e.g., in dromiids) or not (the common feature). Projections on the sternum and abdomen, such as distinct spines that may be present (e.g. portunids) or absent (e.g. pinnotherids).


Megalopae of Dromiidae

Hypoconcha sabulosa (DV)

Dromia erythropus (DV)

Hypoconcha arcuata (DV)

Cryptodromiopsis antillensis (DV)

Fig. 22. Megalopae of Dromiidae. DV: dorsal view; scale in mm. Sources, from Kircher (1970): C; from Lang and Young (1980): A; from Laughlin et al. (1982): B; from Rice and Provenzano (1966): D.

Family and species

Mx1 | Mx2 Mxp2 Mxp3

Carapace outgrowths endopod | endopod exopod exopod

setation I setation setation setation

DROMIIDAE

Cryptodromiopsis antillensis '

; Dromia erythropus ;

! Hypoconcha arcuata

. Hypoconcha sabulosa

LATREILLIIDAE

i Latreillia elegans spines and extremely long 'antlers' 5 j_ 3 0, 6 3^5

with spjnes^hairy^ _ 2 4- |_ 8 10/12_ 4^9 with_spines_ A 2 / _ l ! §_ _Jtli7__ 1<L2 _ with spines 5_ j 4 2, 6 1, 6

anterodorsai spines 2 ,2 ,4 ! 1,5 2,7 0,7

CALAPPIDAE

smooth Hepatus pudibundus

LEUCOSIIDAE

Persephona mediterranea mid- and posterodorsal spines 0

_ 4 _ ; _6, 1

0,3 11

Table 15. Morphological characters based on original descriptions (see Table 14) of the megalopa of seven species among four brachyuran families from the South Atlantic. Setation data refer to numbers of setae per segment, progressing proximally to distally. Mxl: maxillule; Mx2: maxilla; Mxp2: maxilliped 2; Mxp3: maxilliped 3.


Megalopae

Latreilliidae

o Calappidae Parthenopidae Hymenosomatidae

®

Latreillia elegans (DV)

Hepatus Parthenope (Platylambrus) pudibundus (DV) serrata (DV)

Halicarcinus planatus (DV)

Gecarcinidae Atelecyclidae Leucosiidae

Cardisoma guanhumi (DV) Peltarion splnulosum (DV) Persephona mediterranea (DV)

Fig. 23. Megalopae of Latreilliidae, Calappidae, Parthenopidae, Hymenosomatidae, Gecarcinidae, Atelecyclidae, and Leucosiidae. DV: dorsal view; scale in mm: Sources, from Boschi et al. (1969): D; from Costlow and Bookhout (1968): E; from lorio (1983): F; from Negreiros-Fransozo et al (1989): G; from Rice and Williamson (1977): A; from Rieger and Hebling (1993): B; from Yang (1971): C.

Acknowledgements

We thank Dr. Gustavo A. S. de Melo from the Museum of Zoology at the University of Sao Paulo (MZUSP) for providing taxonomic and biogeographical data. Other information was kindly provided by Drs. Nilton J. Hebling from the Universidade Estadual Paulista in

Rio Claro, Brazil, Austin B. Williams from the National Museum of Natural History in Washington, USA, Lucrecia C. Zolessi from the Museum of Natural History in Montevideo, Uruguay, Danilo K. de Calazans from Fundacao Universidade de Rio Grande, Brazil, and Mr. Ignacio Gonzalez-Gordillo from the Institute of Marine Sciences in Andalucia, Spain. Dr.


Richard E. Dowds is thanked for providing illustrations and descriptions of the gennus Pinnixa.

Suggested readings

Boschi E.E. 1981. Larvas de Crustacea Decapoda. In "Atlas del Zooplancton del Atlantico Sudoccidental y metodos de trabajo con el zooplancton marino" (D. Boltovskoy, ed.), Public. Esp. Inst. Nac. Inv. Desarrollo Pesq., Mar del Plata, pp. 699-757. Albeit a somewhat outdated study, it covers non-brachyu-ran decapods and stomatopods from the area that are not included in the present work.

Felder D.L., Martin J.W., Goy J.W. 1985. Patterns in early postlarval development of decapods. In "Larval growth" (A.M. Wenner, ed.), Crustacean Issues Vol. 2, A. Balkema, Rotterdam, pp. 163-225. A good review of decapod developmentafcpatterns.

Gore R.H. 1985. Molting and growth in decapod larvae. In "Larval growth" (A.M. Wenner, ed.), Crustacean Issues Vol. 2, A. Balkema, Rotterdam, pp. 1-66. A good review of decapod larval development that complements Felder et al. (1985).

Gurney R. 1942. The larvae of decapod Crustacea. Ray Soc. Publ., London, pp. 1-306. The classic but still informative review of decapod larvae.

Ingle R.W. 1992. Larval stages of northeastern Atlantic crabs. Natur. Hist. Publ., Chapman & Hall, London, pp. 1-363. An excellent and very detailed account of larvae that includes relevant information on morphology, rearing methods, and a revealing account of the earliest larval studies.

Marques F., Pohle G. 1995. Phylogenetic analysis of the Pinnotheridae (Crustacea: Brachyura) based on larval morphology, with emphasis on the Dissodactylus species complex. Zool. Scripta, 24:347-365. One of very few larval phylogenetic studies using strict cladistic methodology.

Megalopae of Belliidae

Corystoides chilensis (DV)

Acanthocyclus gayi (DV) Acanthocyclus

albatrossis <DV)

Fig. 24. Megalopae of Belliidae. DV: dorsal view; scale in mm. Sources, from Boschi and Scelzo (1970): A; from Campodonico and Guzman (1973): C; from Fagetti and Campodonico (1970): B.


Megalopae of Portunidae

Callinectes sapidus (DV)

Oval ipes catharus (DV)

Portunus spinicarpus (DV)

F ^ . 25. Megalopae of Portunidae. DV: dorsal view; scale in mm. Sources, from Bookhout and Costlow (1974): E; from Costlow and Bookhout (1959): A; from Stuck and Truesdale (1988): B; from Wear and Fielder (1985): C-D (orig. fig. 132 on p. 51).

McConaugha J.R. 1985. Nutrition and larval growth. In "Larval growth" (A.M. Wenner, ed.), Crustacean Issues Vol. 2, A. Balkema, Rotterdam, pp. 127-154. A review and analysis showing the importance of nutrition in growth and development through the larval stages.

Paula J. 1996. A key and bibliography for the identification of zoeal stages of brachyuran crabs (Crustacea, Decapoda, Brachyura) from the Atlantic coast of Europe. J. Plankton Res., 18:17-27. A good guide to the identification of zoeal stages of brachyuran crabs.

Pohle G., Marques F. 1998. Phylogeny of the Pinnotheridae: larval and adult evidence, with emphasis on the evolution of gills. Invertebrate Reproduction and Development, 33:229-239. A

rare phylogenetic study that includes larval and adult evidence in the reconstruction of evolutionary relationships.

Rice A.L. 1980. Crab zoeal morphology and its bearing on the classification of the Brachyura. Trans. Zool. Soc. London, 35:271-424. A classic work and first serious attempt at summarizing the copious larval information since Gurney's (1942) review.

Rice A.L. 1983. Zoeal evidence for brachyuran evolution. In "Crustacean phylogeny" (F.R. Schram, ed.), Crustacean Issues Vol. 1, A. Balkema, Rotterdam, pp. 313-329. A complementary article to the 1980 work that focuses on the evolution of primitive and more advanced groups of crabs based on larval information.


Schmitt W.L. 1971. Crustaceans. Univ. Michigan Press, Ann Arbor, pp. 1-204. An old but very readable and informative text on Crustacea in general.

Scholtz G., Richter S. 1995. Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zool. J. Linn. Soc, 113:289-328. The most recent comprehensive study on the relationships of the major decapod groups.

Warner G.F. 1977. The biology of crabs. Van Nostrand Rheinhold, New York, pp. 1-202. A

good account of many aspects of crab biology, including life histories.

Williamson D.I. 1982. Larval morphology and diversity. In "Biology of Crustacea. Vol. 2. Embryology, morphology, and genetics" (L.G. Abele, ed.), Acad. Press, NewYork, pp. 43-110. The most detailed account of larval morphology and diversity; includes comprehensive keys.

Family and species Rostrum orientation

and morphology

Carapace outgrowths; Mx1 Mx2 Mxp1 Mxp3 or spines on sternal [ endopod endopodj epipod j epipod

seement4 j setation I setation i setation! setation

HYMENOSOMATIDAE

I Halicarcinus pianatus ! directed anteriorly | carapace smooth

ATELECYCLIPAE Pf^lfa nan \

I /rW A , , . . . i deflected ventrally ! carapace with spines I (Ptatylambrusjspinulosuml I

21

25

BELLMDAE

Acanthoc clus albatrossis \

Acanthocyclus gayi


short

short/bilobed

short/bilobed

carapace smooth j 1, 2

carapace with protuberances

2,4

carapace with anterolateraI spines

2,2

PORTUNIDAE

Arenaeus cribrarius directed anteriorly/

developed

Caliinectes sapidus

Ovalipes catharus _


directed anteriorly,

developed deflected ventrally directed anteriorly,

developed

sternal spines present; 2, 2

sternal spines presenti 6, 4 i

sternal spines present ?_

sternal spines present1 6, 2

_1_3_

15

14

26

18

24

18

26

±23

15

±27 Scylia serrata directed anteriorly,

developed sternal spines presentj 3, 2

Table 16. Morphological characters based on original descriptions (see Table 14) of the megalopa often species among four families from the South Atlantic. Setation data refer to numbers of setae per segment, progressing proximally to distally. Abbreviations as in Table 15.


Megalopae of Majidae

Mithrax hispidus (DV)

O Microphrys bicornutus

(DV)

Mithraculus forceps (DV) Libinia

spinosa (DV)


Libidoclaea granaria (DV)

Fig. 26. Megalopae of Majidae. DV: dorsal view; scale in mm. Sources, from Bakker et al. (1990): N; from Bolanos et al. (1990): A; from Bolanos et al. (1996): J; from Boschi and Scelzo (1968): M; from Campodonico and Guzman (1972): I; from Fagetti (1969): P; from Fagetti and Campodonico (1971): G; from Fransozo andHebling (1982): H; from Fransozo and Negreiros-Fransozo (1997): D; from Gore era?. (1982): K; from Fliyodo et al. (1994): F; from Negreiros-Fransozo and Fransozo (1991): C; from Negreiros-Fransozo and Fransozo (1996): Q; from Sandifer and van Engel (1972): E; from Scotto and Gore (1980): O; from Wilson et al. (1979): L; from Yang (1976): B.


Family and species

MAJIDAE

Acanthonyx scutiformis

Anasimus latus





Libinia ferreirae

Libinia spinosa i Microphrys bicornutus

Mithraculus coryphe

Mithraculus forceps

Mithrax caribbaeus

Mithrax hispidus

Pitho Iherminieri

\Pyromaia tuberculata


Taliepus dentatus

PARTHENOPIDAE

1 Parthenope i (Platylambrus) serrata

Rostrum orientation

deflected ventrally

?

deflected ventrally

deflected ventrally

deflected ventrally

directed anteriorly?

deflected ventrally

deflected ventrally

deflected ventrally

directed anteriorly

directed anteriorly

deflected ventrally

directed anteriorly

deflected ventrally

directed anteriorly

deflected ventrally

directed anteriorly?

deflected ventrally

Carapace outgrowths

smooth

spines/protuberances

smooth

smooth

spines

protuberances

protuberances

spines/protuberances

protuberances

protuberances

protuberances

protuberances

protuberances

protuberances

protuberances

sp i nes/protu berances

protuberances

posterodorsal spines

Mx1

endopod

setation ;

1,1

0

0

0

3

1

4

2

0

0

0

0,2

0

0,2

0,2

0,4

3

7 i

Mx2

endopod

setation

0

1

0

0

2

0

2

0

0

0

0

0

2

0

0

3

2

8

Mxp1

epipod

setation

6

4

5

7

12

14

7

4

5

6/7

5

6

5

5

7

8

11

7

Mxp3

epipod

setation

3

3

6

10

2

9

9

6

5

2/8

5

5

3

6

8

5?

9

10

Table 17. Morphological characters based on original descriptions (see Table 14) of the megalopa of 18 species among Majidae and Parthenopidae from the South Atlantic. Setation data refer to numbers of setae per segment, progressing proximally to distally. Abbreviations as in Table 15.


Megalopae of Xanthidae

Hexapanopeus caribbaeus (DV)

Panopeus americanus (DV) Panopeus austrobesus (DV)

Eurypanopeus depressus (DV)

Micropanope sculptipes (DV)

O

Platyxanthus crenulatus (DV)

Menippe nodifrons (DV) Eriphia gonagra (DV)

Hexapanopeus schmitti (DV) Platyxanthus

patagonicus (DV)


Hexapanopeus Panopeus Panopeus paulensis (DV) herbstii (Dv) occidentalis (DV)

Fig. 27. Megalopae of Xanthidae. DV: dorsal view; scale in mm. Sources, from Andryszak and Gore (1981): F; from Bakker et al. (1989): L; from Bookhout and Costlow (1979): P; from Costlow and Bookhout (1961a): G; from Costlow and Bookhout (1961b): R; from Costlow and Bookhout (1966a): O; from Fransozo (1987): K; from Fransozo et al. (1989): Q; from Gore et al. (1981): A; from Ingle (1985): S; from lorio and Boschi (1986): M; from Kurata et al. (1981): N; from Martin et al. (1985): I; from Memi-Marque (1970): H; from Montii et al. (1988): E; from Negreiros-Fransozo (1986a): B; from Negreiros-Fransozo (1986b): D; from Scotto (1979): J; from Vieira and Rieger (in press): C.


Species Rostrum orientation

and morphology

Carapace

outgrowths

Mx1 Mx2 Mxp1 I Mxp3

endopod! endopod epipod i

setation ! setation I setation

epipod

setation

XANTHIDAE

Eriphia gonagra

Eurypanopeus abbreviatus

E. depressus

Eurytium iimosum

Garthiope barbadensis

Hexapanopeus angustifrons

H. caribbaeus

H. pauiensis

H. schmitti

Menippe nodifrons

Micropanope sculptipes

Panopeus americanus

P. austrobesus

P. bermudensis

P. herbstii

P. occidentalis

Pilumnus dasypodus

Platyxanthus crenulatus

P. patagonicus

short, deflected ventrally

short, deflected ventral ly

short/ deflected ventrally

directed anteriorly





short, directed anteriorly


deflected ventrally

short


short

short



short, bifid


smooth

smooth

protuberances

protuberances

smooth

smooth

smooth

smooth

smooth

smooth

protuberances

smooth

smooth

smooth

smooth

smooth

smooth

smooth

smooth

3,4

3

1,6

?

5

7

1,2

4

2,3

1,4

?

2,3

1,4

2 3

4,3

5 1 1(2), 4(6) I

| 4 1 4

5

4

8

?

5

7

2

2

2

4-6

?

1

8(9)

1

7

1(2)

9

9

6

18

5

7

?

7

7

6

8

9

12-26

?

7

6

5

7

1(2)

5

10

I 7

38

16

12

?

15

± 2 0

20

18 \

13-15

26-36 •

?

12 |

6 !

16

20

6(7)

11(13)

20

20

Table 18. Morphological characters based on original descriptions (see Table 14) of the megalopa of 19 species of Xanthidae from the South Atlantic. Setation data refer to numbers of setae per segment, progressing proximally to distally. Abbreviations as in Table 15.

Megalopae of Pinnotheridae

Clypeasterophilus stebbingi (DV)

Tumidotheres maculatus (DV)

Dissodactylus crinitichelis (DV)

Fig. 28. Megalopae of Pinnotheridae. DV: dorsal view; scale in mm. Sources, from Costlow and Bookhout (1966b): B; from Marques and Pohle (1996b): A; from Pohle and Telford (1981): C.


Family and species

PINNOTHERIDAE


Dissodactylus criniticheiis

Tumidotheres maculatus

GRAPSIDAE

Aratus pisonii

Armases angustipes

Chasmagnathus granufata

Cyclograpsus integer

Cyrtograpsus angulatus

C. aftimanus

Metasesarma rubripes

Percnon gibbesi

Sesarma rectum

GECARCINIDAE

Cardisoma guanhumi

OCYPODIDAE

Ocypode quadrata

Uca burgersi

U. mordax

U. thayeri

U. uruguayensis

U. vocator

Ucides cordatus

Rostrum orientation

and morphology

directed anteriorly

absent

short

absent

short

absent

deflected ventrally

short, deflected ventral ly

deflected ventrally

short

trifid

short

deflected ventrally

deflected ventrally

deflected ventrally

deflected ventrally

deflected ventrally

deflected ventrally

deflected ventrally

deflected ventrally

Carapace

outgrowths

smooth

smooth

three long spines

dorsal spine

smooth

smooth

smooth

smooth

smooth

smooth '

smooth

protuberances

smooth

smooth

smooth

smooth

smooth

smooth

smooth

smooth

M x l

endopod

setation

0,4

2,1

1,5

3,4

5

6

1,3

6

2,3

?

1,5

2 ,4

3

1,2

1,1

^ 0 , 4

2

0,1

0,5

Mx2

endopod

setation

2

3

0

5

0

0

2

0

2

0

?

0

6

7

2

0

0

0

2

6

Mxp1

epipod

setation

4

4

4

?

7

7

9

7

8

7

?

5

9

± 2 0

7

6

7

6(7)

5

10

Mxp3

epipod

setation

26?

16

21 ?

?

16

16

± 3 5

22

35

20

?

15

25

± 5 4

17

19

16

± 1 9

13

32

Table 19. Morphological characters based on original descriptions (Table 14) of the megalopae of 20 species among 4 families from the South Atlantic. Setation data refers to numbers of setae per segment, progressing proximally to distally. Abbreviations as in Table 15.


Megalopae of Grapsidae

Cyrtograpsus angulatus (DV)

Armases angustipes (DV)


Ucides cordatus (DV)

Ocypode quaarata (DV)

Megalopae of Ocypodidae

Uca burgersl (DV)

Uca thayeri (DV)

Fig. 30. Megalopae of Ocypodidae. DV: dorsal view; scale in mm. Sources, from Anger et al. (1990): G; from Diaz and Costlow (1972): D; from Rieger (1992): C, F; from Rieger (1996): from Rieger (1998): E; B; from Rodrigues and Hebling (1989): A.

Fig. 29. Megalopae of Grapsidae. DV: dorsal view; scale in mm. Sources, from Boschi et al. (1967): D; from Diaz and Ewald (1968): E (orig. fig. 5 on p. 235); from Fransozo and Hebling (1986): G; from Gore and Scotto (1982): B; from Kowalczuk (1994): I; from Paula and Hartnoll (1989): C; from Rieger and Vieira (1997): H; from Scelzo and Lichstein de Bastida (1979): A; from Warner (1968): F.

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